Professional Documents
Culture Documents
Lim
Edible Medicinal
and Non-Medicinal
Plants
Volume 11,
Modified Stems, Roots, Bulbs
Edible Medicinal and Non-Medicinal
Plants
T.K. Lim
Special thanks for the use of digital images are accorded to Cecilia Lafosse
(CIP) and Ezeta Fernando (ex CIP), International Potato Centre (CIP) for oca
(Oxalis tuberosa). Frank Van Keirsbilck for mauka (Mirabilis expansa) and
GF Chung for Indian goosegrass (Eleusine indica).
v
Disclaimer
The author and publisher of this work have checked with sources believed to
be reliable in their efforts to confirm the accuracy and completeness of the
information presented herein and that the information is in accordance with
the standard practices accepted at the time of publication. However, neither
the author nor publishers warrant that information is in every aspect accurate
and complete and they are not responsible for errors or omissions or for con-
sequences from the application of the information in this work. This book is
a work of reference and is not intended to supply nutritive or medical advice
to any individual. The information contained in the notes on edibility, uses,
nutritive values, medicinal attributes and medicinal uses and suchlike included
here are recorded information and do not constitute recommendations. No
responsibility will be taken for readers’ own actions.
vii
Contents
Introduction ........................................................................................... 1
Iridaceae
Lamiaceae
Marantaceae
Nelumbonaceae
Nyctaginaceae
Nymphaeaceae
Orchidaceae
ix
x Contents
Oxalidaceae
Piperaceae
Poaceae
Rubiaceae
Simaroubaceae
This book continues as volume eleven of a multi- edible fruits or flowers have been covered in ear-
compendium on Edible Medicinal and Non- lier volumes and for those better known for other
Medicinal Plants. It covers plants with edible non-reproductive plant parts will be covered in
modified storage subterranean stems (corms, rhi- latter volumes.
zomes, stem tubers) and unmodifed subterranean As in the preceding ten volumes, topics cov-
stem stolons, above-ground swollen stems and ered include: taxonomy (botanical name and syn-
hypocotyls, storage roots (tap root, lateral roots, onyms); common English and vernacular names;
root tubers) and bulbs that are eaten as conven- origin and distribution; agro-ecological require-
tional or functional food as vegetables and spices, ments; edible plant part and uses; plant botany;
as herbal teas, and may provide a source of food nutritive and medicinal/pharmacological proper-
additive or neutraceuticals. A list of such edible ties with up-to-date research findings;traditional
plant species from families Acanthaceae to medicinal uses; other non-edible uses; and
Zygophyllaceae are presented in a tabular form selected/cited references for further reading.
and 32 such edible species from the families A corm or bulbotuber is defined as a short,
Alismataceae, Amaryllidaceae, Apiaceae, vertical, swollen underground plant stem that
Araceae, Araliaceae, Asparagaceae, Asteraceae, serves as a storage organ used by some plants to
Basellaceae, Brassicaceae and Campanulaceae survive unfavourable adverse periods. It bears
had been covered in detail in preceding volume 9. membranous or scaly leaves and buds. Some
Nineteen edible species from the families examples of plants with edible corms are found
Amaranthaceae, Cannaceae, Cibotiaceae, in Amorphophallus spp., Colocasia esculenta
Convolvulaceae, Cyperaceae, Dioscoreaceae, (taro), Eleocharis dulcis (Chinese water chest-
Euphorbiaceae and Fabaceae had been covered in nut), Sagittaria spp. (arrowhead or wapato) and
detail in volume 10. This volume 11 covers in Xanthosoma spp. (cocoyam or tannia). Corms
detail 18 edible species in the families Iridaceae often give rise to many small secondary corms or
(1), Lamiaceae (1), Marantaceae (1), cormlet called cormels at the end of very short
Nelumbonaceae (1), Nyctaginaceae (1), stolons.
Nymphaeaceae (3), Orchidaceae (4), Oxalidaceae Rhizome is a modified subterranean stem of a
(1), Piperaceae (1), Poaceae (2), Rubiaceae (1) plant that is usually found underground, produc-
and Simaroubaceae (1). Other species from these ing roots and shoots. It is used by the plant as
families with edible modified stems, roots and storage organ and whole rhizome or pieces of the
bulbs are listed in Table 1. Many plants with such rhizome serves as vegetative propagules to give
edible plant parts that are better known for their rise to new plants. Examples of plants with edible
rhizomes include gingers (Zingiber spp.), tur- leaves or leaf bases that serve as food storage
meric (Cucurma longa), greater galangal (Alpinia organs during dormancy and enable the plant to
galanga), lesser galangal Alpinia officinarum), survive through adverse periods. The fleshy
sand ginger or kencur (Kampferia galanga), lotus leaves are arranged in a concentric manner. Bulbs
root (Nelumbo nucifera), Typha spp., fingerroot can be tunicate i.e. with membranous papery cov-
(Boesenbergia rotunda) and arrowroot (Maranta ering (scale leaves) or tunic that protects the inner
arundinacea). fleshy scale leaves from drying and mechanical
A stem tuber is a modified plant storage organ injury. Examples of tunicate bulbs are the
that is formed from thickened rhizome or stolon. Alliums, onions, leeks, hyacinth and tulips. In
The tops or sides of the tuber produce shoots that imbricate or non-tunicate bulbs, the fleshy scale
grow into typical stems and leaves and the under- leaves are not in concentric rings but are loosely
sides produce roots. The stem tuber has all the arranged or spreading, overlapping one another
parts of a normal stem, including nodes (eyes) and at the margin. Such a bulb is not a compact body
internodes. A stem tuber may start off as an and not usually covered by a common tunic.
enlargement of the hypocotyls of the seedling and Examples are the garlic (Allium sativum) and
may include the epicotyl or upper section of the some Lilium lilies.
root as is in the case of maca (Lepidium meyenni). Tap root is the true main root of the plant and in
More commonly as in Plectranthus esculenta in some species the tap root is modified and fleshy,
the Lamiaceae family, numerous tubers are rich in stored nutrients; they may or may not be
formed on short stolons that arise from the base of fused with the hypocotyl or basal stem tissues and
the stem, or as in potatoes tubers are formed as maybe napiform, globose, conical, fusiform or
enlarged stolons thickened and enlarged into stor- cylindrical in shape. Notable examples of plants
age organs. In some Cyperus species e.g. tigernut with edible tap roots are Abelmochus spp., beet
or chufa (C. esculentus), the stolons end with the (Beta vulgaris), rutabaga, turnip, Bunium persi-
growth of tubers that can give rise to new plants. cum, burdock, carrot, radish and daikon, celeriac,
Other striking examples of plants with stem tubers jicama and ahipa (Pachyrhizus spp.), parsnips,
include hog potato or groundnut (Apios ameri- parsley, skirret (Sium sisarum), bush potato (Vigna
cana), Jerusalem artichoke or sunchoke lanceolata), salsify (Tragopogon porrifolius),
(Helianthus tuberosus), earthnut pea (Lathyrus black salsify (Scorzonera hispanica), tongkat Ali
tuberosus), oca or New Zealand yam (Oxalis (Eurycoma longifolia) and many others. Plants
tuberosa), Chinese artichoke or crosne (Stachys with edible root tubers or tuberous roots with
affinis), mashua or añu (Tropaeolum tuberosum) enlarged root and lateral roots function as storage
and ulluco (Ullucus tuberosus). In Botany, a sto- organs, lacking nodes, internodes and adventious
lon is an horizontal modified stem arising from buds. Notable examples include pignut or earthnut
the base of a plant that produces new plants from (Conopodium majus), sweet potato (Ipomoea bata-
buds at its tip or nodes and forms adventitious tas), desert yam (Ipomoea costata), cassava or yuca
roots at the nodes, it can be creeping above the or manioc (Manihot esculenta), yams (Dioscorea
ground surface or underground. An example of a spp.), mauka or chago (Mirabilis expansa), bread-
plant with edible stolon is Imperata cylindrica. root, tipsin, or prairie turnip (Psoralea esculenta)
However, some botanists used the term stolons for and yacón (Smallanthus sonchifolius).
stem branches that arise from the base of the stem Most terrestrial orchids are rhizomatous or
that creeps above the ground and those that creeps have corms or tubers. Many orchid species pro-
horizontally underground as rhizomes. An exam- duce edible tubers and roots while comparatively
ple of a plant with swollen, above-ground storage fewer species (e.g. Cymbidium canaliculatum
stem is the kohlrabi. and Dendrobium tarberi) produce edible pseudo-
Bulb is a much reduced underground stem bulbs. These tubers contain a nutritious, starchy
bearing at its apex a growing or floral primor- polysaccharide called glucomannan. These
dium surrounded by thick, fleshy modified scale tubers provide a starchy flour called salep which
Introduction 3
Table 1 Plants with edible modified stems, roots and bulbs in the families: Iridaceae, Lamiaceae, Marantaceae,
Nelumbonaceae, Nyctaginaceae, Nymphaeaceae, Orchidaceae, Oxalidaceae, Piperaceae, Poaceae, Rubiaceae and
Simaroubaceae
Common/Vernacular
Family Scientific Name Names Edible Part Use Reference
Iridaceae Crocus sativus L. Saffron Roots are eaten roasted Hedrick (1972),
Kunkel (1984),
Morton (1976),
Facciola (1990),
and Lim (2014)
Iridaceae Iris cristata Aiton Dwarf Crested Iris Roots used as a spice. Hedrick (1972),
Frequently chewed by Tanaka (1976),
local people to alleviate Kunkel (1984),
thirst. When first and Coffey (1994)
chewed, the roots have
a pleasant sweet taste,
within a few minutes
this changes to a
burning sensation far
more pungent than
capsicums
Iridaceae Iris x germanica L. Bearded Iris, Flag, Orris Root used as spice, Chase (1900),
Root, German Iris, flavouring in ice cream, Parmentier (1781)
Florentine Iris confectionery and cited by
baked goods; starch Freedman (2009),
used as bread flour Uphof (1968),
Morton (1976),
Facciola (1990),
Bender (2009),
and Surhone et al.
(2011)
Iridaceae Iris pallida Lam. Sweet Iris, Dalmation Orris oil from rhizome Morton (1976)
Iris used to flavour soft
drinks, candy and
chewing gum
Iridaceae Iris setosa Pall. ex Link Hiogi-Ayame Rhizomes eaten or used Uphof (1968),
as source of starch Hedrick (1972),
Tanaka (1976),
and Facciola
(1990)
Iridaceae Morea fugax Uintjie Bulbous root eaten Hedrick (1972),
(D. Delaroche) Jacq. roasted, boiled or Fox et al. (1982),
stewed with milk and Facciola
(1990)
Iridaceae Romulea bulbocodium NF Bulbous root eaten Fairchild (1930)
(L.) Sebast. & Mauri
Iridaceae Tigrida pavonia (L. f.) Common Tiger Flower, Roasted starchy corms Uphof (1968) and
Redouté Jockey Cap, Mexican used as food by Facciola (1990)
Shellflower, Peacock Mazatecs and other
Flower Indian tribes in Mexico
Lamiaceae Callicarpa rubella Gopura Esing (Mishing) Bark and roots chewed Patiri and Borah
Lindl. Bonmala (Assamese) like betel nut; roots (2007) and
eaten in Meghalaya Sawian et al.
(2007)
(continued)
4 Introduction
Table 1 (continued)
Common/Vernacular
Family Scientific Name Names Edible Part Use Reference
Lamiaceae Callicarpa vestita Wall. Yarpo Esing (Mishing) Bark and roots chewed Patiri and Borah
ex C.B.Clarke like betel nut (2007)
Lamiaceae Clerodendrum fragrans Fragrant Glorybower; Roots dried, cooked Hu (2005)
(Vent.) R.Br. = Chou Mo Li, Chou Mu with pork to strengthen
Clerodendrum chinense Dan (Chinese) elderly people and to
(Osbeck) Mabb, remove pain and
stiffness of muscles and
joints
Lamiaceae Clerodendrum serratum Phelang Riho Roots eaten in Karbi, Kar and
(L.) Moon = Rotheca (Assamese) Assam Borthakur (2008)
serrata (L.) Steane &
Mabb.
Lamiaceae Coleus blumei Benth. = Coleus, Painted Nettle, Tubers eaten Burkill (1966)
Plectranthus Sayabana, Jacob’s Coat and Facciola
scutellarioides (L.) (1990)
R.Br.
Lamiaceae Coleus dazo A. Chev. = Daju, Rizuka Starchy root peeled, Tanaka (1976)
Plectranthus esculentus boiled, served and and Facciola
N.E.Br. eaten or pickled (1990)
Lamiaceae Coleus parviflorus African Potato, Country Tubers eaten like Tanaka (1976),
Benth. = Plectranthus Potato potatoes Ochse and van
rotundifolius (Poir.) den Brink (1980)
Spreng. and Facciola
(1990)
Lamiaceae Coleus tuberosus African Potato, Country Tubers usually eaten Ochse and van
(Blume) Benth. = Potato steamed or cooked with den Brink (1980)
Plectranthus rice In Indonesia
rotundifolius (Poir.)
Spreng.
Lamiaceae Eriophyton wallichii Mian Shen (Chinese) Roots used in food in Hu (2005)
Benth. ex Wall. north-western Yunnan
Lamiaceae Leonurus sibiricus L. Siberian Motherwort In China, the roots are Burkill (1966),
cooked with pork Altschul (1973),
Tanaka (1976),
and Facciola
(1990)
Lamiaceae Lycopus europaeus L. Gypsywort, Water China: root eaten. Read (1946) and
Horehound Manchuria: starchy Baranov (1967)
tubers eaten
Lamiaceae Lycopus lucidus Turcz. Lycopos, Bugleweed; Di Underground rhizomes Hu (2005)
ex benth. Gua Er Miao (Chinese) eaten in northern china
and Yunnan
Lamiaceae Lycopus uniflorus Northern Bugleweed White tubers eaten raw Fernald et al.
Michx. in salads, boiled, (1985) and
pickled or added to Facciola (1990)
soups and stews
Lamiaceae Phlomis tuberosa L. Tuberous Jerusalem Roots eaten by the Hedrick (1972)
Sage; Bodmon Sok Kalmucks in Eurasia and Facciola
(1990)
Lamiaceae Plectranthus barbatus Indian Coleus Tubers eaten Jansen (1996)
Andrews
(continued)
Introduction 5
Table 1 (continued)
Common/Vernacular
Family Scientific Name Names Edible Part Use Reference
Lamiaceae Plectranthus edulis Galla Potato Tubers eaten Jansen (1996)
Agnew
Lamiaceae Plectranthus esculentus Livinstone Potato, Kaffir Stem tubers eaten, Jansen (1996),
N.E. Br. Potato eaten raw or boiled and Phillips and Rix
eaten as vegetables (1993), van Wyk
(2006), and
Codex (2014)
Lamiaceae Plectranthus Madagascar Spur Flower Tubers eaten in Tanaka (1976)
madagascariensis Madagascar and Facciola
(Pers.) Benth. (1990)
Lamiaceae Plectranthus Chinese Potato, Coleus Young aromatic tubers Jansen (1996) and
rotundifolius (Poir.) Potato, Hausa Potato used in soup and Codex (2014)
Spreng vegetable dishes
Lamiaceae Solenostemon Huasa Potato, Fra-Fra Nigeria (Kano State, Dalziel (1955),
rotundifolia (Poir.) Potato; Tumuku, Tamaka northern): tuber eaten Fox et al. (1982),
J.K. Morton. = (Hausa) like potato Mortimore
Plectranthus (1989), Facciola
rotundifolius (Poir.) (1990), and
Spreng Codex (2014)
Lamiaceae Stachys adulterina Hubei Artichoke; Di Can Root tubers used as Hu (2005)
Hemsl. Zi (Chinese) vegetables, cooked or
pickled
Lamiaceae Stachys affinis Bunge Chinese Artichoke; Cao As above Phillips and Rix
Shi Can (Chinese) (1993) and Hu
(2005), Codex
2014
Lamiaceae Stachys chinensis Bunge Hyssopleaf Hedgenettle Manchuria: rhizome Baranov (1967)
ex Benth eaten
Lamiaceae Stachys sieboldi Miq. Crosnes, Chinese Japan: tubers salted or Read (1946),
Artichoke, Japanese preserved in plum Facciola (1990),
Artichoke vinegar Van den Bergh
(1996), and
Codex (2014)
Marantaceae Calathea allouia Guinea Arrowroot, Root tubers boiled and Facciola (1990),
(Aublet) Lindl. Leren, Sweet Corn Tuber eaten like potato Groen et al.
(1996), and
Codex (2014)
Marantaceae Halopegia blumei Jelantir (Javanese), Patat Tubers eaten cooked or Groen et al.
(Körn) K Schum. (Sundanese) Dong Nam roasted (1996), Ochse
(Vietnam) and van den Brink
(1980), and
Ochse and van
den Brink (1980)
Marantaceae Maranta arundinacea L. Arrowrot, Tora Alu, Tha Rhizomes are source of Ochse and van
Lairusa, Hnathel, arrowroot, eaten den Brink (1980),
Hpogimbai (Assamese) cooked or raw Tuber Facciola (1990),
Khaita Alu (Boro) eaten both raw and Villamayor Jr and
Nginti Ali (Mishing) boiled, starch from Jukema (1996),
rhizome Hu (2005), Patiri
and Borah (2007),
Medhi and
Borthakur (2012),
and Codex (2014)
(continued)
6 Introduction
Table 1 (continued)
Common/Vernacular
Family Scientific Name Names Edible Part Use Reference
Marantaceae Maranta dichotoma Mohtra Reed, Sitalpati Tuber eaten both raw Medhi and
(Roxb.) Wall. = Plant; Tha Lairu, and boiled Borthakur (2012)
Schumannianthus Hnathel, Hpogimbai
dichotomus (Roxb.) (Assamese)
Gagnep.
Marantaceae Phrynium capitatum Packing Leaf Root tuber eaten in Sawian et al.
Willd. = Phrynium Meghalaya (2007)
pubinerve Blume
Marantaceae Thalia geniculata L. Swamp Lily Rhizomes baked and Tanaka (1976)
eaten or made into a and Facciola
kind of arrowroot (1990)
Nelumbonaceae Nelumbium speciosum Pink Water Lily India: root and seeds Gammie (1902)
Willd. eaten and Watt (1908)
Nelumbonaceae Nelumbo lutea Pers. American Lotus, Water Large tubers, when Facciola (1990)
Chinquapin baked, are sweet and and Saunders
mealy with a flavour (1920)
somewhat like a
sweetpotato
Nelumbonaceae Nelumbo nucifera Lotus, Lotus Root Root eaten raw or Burkill (1966),
Gaertn. cooked, sliced used in Cribb and Cribb
stir-fries, soups, stews (1987), Facciola
or fried as a garnish or (1990), Phillips
side dish. Sliced pieces and Rix (1993),
can be candied or Ong (1996), Hu
pickled. Lotus root (2005), Santich
flour is starch and can et al. (2008), van
be used to make Wyk (2006), and
desserts Codex (2014)
Nyctaginaceae Abronia latifolia Yellow Sand Verbena Root edible Yanovsky (1936)
Eschsch. and Facciola
(1990)
Nyctaginaceae Boerhavia coccinea Tar Vine, Hog Weed Bland fibrous tap root Cribb and Cribb
Mill. eaten (1987) and
Harden (1990)
Nyctaginaceae Boerhavia diffusa L. Hog Weed, Horse Fleshy portion of thick Hu (2005)
Purslane; Zhu Er Yan, roots roasted and eaten,
Huang Xi Xin (Chinese) sweetish and nutritious
Nyctaginaceae Boerhavia spp. Tar Vines Bland fibrous tap root Low (1991)
eaten
Nyctaginaceae Mirabilis expansa (Ruiz Mauka, Chago Tuber dried, boiled or Tanaka (1976),
& Pav.) Standl. fried, eaten as Popenoe et al.
vegetable (1989), Facciola
(1990), Bermejo
and Leon (1994),
and Codex (2014)
Nymphaeaceae Euryale ferox Salisb. Chicken Head, Fos Nut In China, roots and Read (1946) and
seeds eaten Facciola (1990)
Nymphaeaceae Nuphar advena R.Br. Common Spatterdock Rootstock eaten raw, Uphof (1968),
roasted or cooked with Hedrick (1972),
meat and Facciola
(1990)
(continued)
Introduction 7
Table 1 (continued)
Common/Vernacular
Family Scientific Name Names Edible Part Use Reference
Nymphaeaceae Nuphar luteum Sibth. & Yellow Water Lily Rootstock boiled as Hedrick (1972),
Sm. vegetable Fernald et al.
(1985), and
Facciola (1990)
Nymphaeaceae Nuphar polysepala Cow Lily, Spatterdock, Rich starchy rhizome Schofield (2003)
Engelm. = Nuphar lutea Pond Collard used as survival food,
subsp. polysepala after boiling, roasting
(Engelm.) E.O. Beal. or baked and skin
removed
Nymphaeaceae Nuphar pumilum Yellow Pond Lily; Ping Young tender rhizomes Hu (2005)
(Timm.) DC. Peng Cao (Chinese) used as potherb in
Yunnan and Hubei
Nymphaeaceae Nymphaea lotus L. Egyptian Lotus, White Tubers edible Tanaka (1976),
Lotus Kunkel (1984),
and Facciola
(1990)
Nymphaeaceae Nymphaea stellata Blue Lotus Of India Rhizomes eaten raw or Uphof (1968),
Willd. = Nymphaea roasted Hedrick (1972),
nouchali Burm.f. Tanaka (1976),
and Facciola
(1990)
Nymphaeaceae Nymphaea alba L. White Lotus, European In France, root Parmentier (1781)
White Waterlily recommended as a (cited by
famine food after Freedman (2009))
cooking in water and
being flavoured
Nymphaeaceae Nymphaea caerulea Blue Lotus Of Egypt, Starchy tubers eaten Tanaka (1976),
Savigny = Nymphaea Blue Water Lily boiled or roasted Fox et al. (1982),
nouchali var. caerulea and Facciola
(Savigny) Verdc. (1990)
Nymphaeaceae Nymphaea edulis DC. Red Water Lily; In India (Madras Shortt
Shunguneer Pushpum Presidency), roots and (1887–1888)
(Tamil); Koteka, seeds cooked and eaten
Kalharamu (Telugu)
Nymphaeaceae Nymphaea gigantea Giant Water Lily Tuberous rootstock Cribb and Cribb
Hook. eaten (1987)
Nymphaeaceae Nymphaea lotus var. Red Water Lily Root eaten baked or Paton and Dunlop
pubescens (Willd) boiled with salt added (1904)
Hook.f. &
Thomson = Nymphaea
pubescens Willd.
Nymphaeaceae Nymphaea lotus L. Egyptian Lotus, White In upper Guinea Africa, Gammie (1902),
Lotus; Bado (Hausa); root used as a famine Watt (1908),
Dambi (Kanuri) food, being either Irvine (1952),
roasted in ashes or Uphof (1968),
dried before being Hedrick (1972),
ground into flour. In Tanaka (1976),
Nigeria (Kano State, Pongpangan and
northern), rhizome and Poobrasert
seeds eaten. In India (1985),
(Bombay Presidency), Mortimore
roots and seeds eaten (1989), and
Facciola (1990)
(continued)
8 Introduction
Table 1 (continued)
Common/Vernacular
Family Scientific Name Names Edible Part Use Reference
Nymphaeaceae Nymphaea nouchali Boga Bhet, Seluk Rhizomes/roots eaten Van den burgh
Burm.f. (Assamese) raw or cooked as (1994) and Patiri
vegetable in Assam and Borah (2007)
Nymphaeaceae Nymphaea odorata Frgarny Water Lily Tubers edible Tanaka (1976),
Aiton Kunkel (1984),
and Facciola
(1990)
Nymphaeaceae Nymphaea rubra Roxb. Ronga Bhet, Mokua, As above Patiri and Borah
ex Andrew Seluk (Assamese) (2007)
Nymphaeaceae Nymphaea spp. Waterlilies Fibrous tubers edible Cribb and Cribb
(1987) and Low
1989
Nymphaeaceae Nymphaea stellata Izibo (Zulu) In Zululand (Ubombo Hely- Hutchinson
Willd. = Nymphaea district), tuber boiled (1898), Gammie
nouchali Burm.f. and eaten. In India, (1902), Watt
roots and seeds eaten (1908) and Uphof
(1968)
Nymphaeaceae Nymphaea tetragona Four Angled Water Lily; Rhizomes used as food Hu (2005)
Georgi Shui Lian (Chinese) in north-west China
Nymphaeaceae Nymphaea tuberosa Tuberous Waterlily, Tubers occasionally Gibbons and
Paine = Nymphaea White Water Lily eaten Tucker (1979),
odorata subsp. tuberosa Fernald et al.
(Paine) Wiersema & (1985), and
Hellq. Facciola (1990)
Nymphaeaceae Ondinea purpurea NF Corms eaten by Les (2003)
Hartog Australian aborigines
Orchidaceae Aceras antropophorum Man Orchid Starchy tubers used for Sezik (2002)
(L.) R.Br. = Orchis making salep flour used
anthropophora (L.) All. in ice-cream
production,
confectionery and
nutritious beverages
Orchidaceae Acianthus apprimus Mountain Gnat Orchid Tubers edible Steenbeeke
D.L. Jones (2001)
Orchidaceae Acianthus collinus NF Tubers edible Steenbeeke
D.L.Jones (2001)
Orchidaceae Acianthus exsertus R.Br Mosquito Orchid Tubers edible Steenbeeke
(2001)
Orchidaceae Acianthus pusillus Gnat Orchid Tubers edible Steenbeeke
D.L. Jones (2001)
Orchidaceae Anacamptis coriophora NF Starchy tubers used for Ghorbani et al.
(L.) R.M.Bateman making salep flour used (2014) and
in ice-cream Kreziou et al.
production, (2015)
confectionery and
nutritious beverages
Orchidaceae Anacamptis morio (L.) Green-Winged Orchid, As above Kreziou et al.
R.M.Bateman Green-Veined Orchid (2015)
(continued)
Introduction 9
Table 1 (continued)
Common/Vernacular
Family Scientific Name Names Edible Part Use Reference
Orchidaceae Anacamptis morio As above As above Ghorbani et al.
subsp. picta (Loisel.) (2014)
Jacquet & Scappat
Orchidaceae Anacamptis palustris NF As above Ghorbani et al.
(Jacq.) R.M.Bateman, (2014)
Pridgeon & Chase
Orchidaceae Anacamptis Butterfly Orchid As above Kreziou et al.
papilionacea (L.) (2015)
R.M.Bateman
Orchidaceae Anacamptis pyramidalis Pyramidal Orchid As above Sezik (1967),
(L.) Rich. Sezik and Özer
(1983) cited by
Tekinşen and
Güner (2010),
Sezik (2002),
Ghorbani et al.
(2014), and
Kreziou et al.
(2015)
Orchidaceae Arthrochilus huntianus Elbow Orchid Tubers probably edible Steenbeeke
(F.Muell.) Blaxell (2001)
Orchidaceae Barlia robertiana Giant Orchid, Orkida Starchy tubers used for Sezik (2002)
(Loisel.) Greuter Kbira making salep flour used
in ice-cream
production,
confectionery and
nutritious beverages
Ochidaceae Brachycorythis Likos, Ligosi As above Mapunda (2007)
pleistophylla Rchb.f. and Hamisy
(2010)
Orchidaceae Calendenia caerula Blue Fairy Orchid, Blue Tubers edible Steenbeeke
R.Br. Caladenia (2001)
Orchidaceae Calendenia carnea Pink Fairies, Pink Fringe Tubers edible Steenbeeke
R.Br. Orchid (2001)
Orchidaceae Calendenia filamentosa Daddy Longlegs, Spider Tubers edible Steenbeeke
R.Br. Orchid (2001)
Orchidaceae Calendenia fuscata Dusky Fingers Tubers edible Steenbeeke
(Rchb.f.) M.A.Clem. & (2001)
D.L.Jones
Orchidaceae Calendenia gracilis Musky Caladenia, Tubers edible Steenbeeke
R.Br. Musky Finger Orchid (2001)
Orchidaceae Calendenia quadrifaria NF Tubers edible Steenbeeke
(R.S.Rogers) D.L.Jones (2001)
Orchidaceae Calendenia tentaculata Green Combed Spider Tubers edible Steenbeeke
Schltdl. Orchid, Fringed Spider (2001)
Orchid
Orchidaceae Chiloglottis diphylla Common Ant Orchid Tubers edible Steenbeeke
R.Br. (2001)
Orchidaceae Chiloglottis formicifera Ant Orchid Tubers edible Steenbeeke
Fitzg. (2001)
(continued)
10 Introduction
Table 1 (continued)
Common/Vernacular
Family Scientific Name Names Edible Part Use Reference
Orchidaceae Chiloglottis palachila Clubbed Ant Orchid Tubers edible Steenbeeke
D.L. Jones & (2001)
M.A. Clem.
Orchidaceae Chiloglottis platyptera NF Tubers edible Steenbeeke
D.L.Jones (2001)
Orchidaceae Chiloglottis pluricallata Brown Bird Orchid Tubers edible Steenbeeke
D.L.Jones (2001)
Orchidaceae Chiloglottis sphyrnoides Ornate Ant Orchid Tubers edible Steenbeeke
D.L.Jones (2001)
Orchidaceae Chiloglottis Broad-Lip Bird Orchid Tubers edible Steenbeeke
trapeziformis Fitzg. (2001)
Orchidaceae Chiloglottis trilabra NF Tubers edible Steenbeeke
Fitzg. (2001)
Orchidaceae Coelogyne ovalis Lindl. Oval Coelogyne Sotirov (2015)
Orchidaceae Comperia comperiana Komper’s Orchid Starchy tubers used for Sezik (2002)
(Steven) Asch. & making salep flour used
Graebn. = in ice-cream
Himantoglossum production,
comperianum (Steven) confectionery and
P.Delforge nutritious beverages
Orchidaceae Corybas fimbriatus Fringed Helmut Orchid Tubers edible Steenbeeke
(R.Br.) Rchb.f. (2001)
Orchidaceae Corybas hispidus Bristly Helmut Orchid Tubers edible Steenbeeke
D.L.Jones (2001)
Orchidaceae Corybas montanus Mt Maroon Helmut Tubers edible Steenbeeke
D.L.Jones Orchid (2001)
Orchidaceae Corybas species A Sphagnum Helmut Tubers edible Steenbeeke
Orchid (2001)
Orchidaceae Cryptostylis erecta Tartan Tongue Orchid, Fleshy starchy root Steenbeeke
F. Muell. ex Benth. Turban Orchid, Bonnet eaten (2001)
Orchid
Orchidaceae Cryptostylis humteriana Leafless Tongue Orchid Fleshy starchy root Steenbeeke
Nicholls eaten (2001)
Orchidaceae Cryptostylis leptochila Small Tongue Orchid, Fleshy starchy root Steenbeeke
R.Br. Red Tongue Orchid eaten (2001)
Orchidaceae Cryptostylis subulata Large Tongue Orchid, Fleshy starchy root Steenbeeke
(Labill.) Rchb.f. Cowslip Orchid eaten (2001)
Orchidaceae Cymbidium Tiger Orchid, Channelled Starch rich pseudobulbs Steenbeeke
canaliculatum R.Br. Cymbidium, Inland Tree eaten cooked or raw (2001)
Orchid
Orchidaceae Dactylorhiza incarnata Early Marsh Orchid Starchy tubers used for Ghorbani et al.
(L.) Soó making salep flour used (2014)
in ice-cream
production,
confectionery and
nutritious beverages
Orchidaceae Dactylorhiza osmanica The Ottoman Starchy tubers used for Tekinşen and
var. Dactylorhiza making salep flour used Güner (2010) and
osmanica = Dactylorhiza in ice-cream Citil and Tekinşen
osmanica (Klinge) P.F production, (2011)
Hunt & Summerh. confectionery and
nutritious beverages
(continued)
Introduction 11
Table 1 (continued)
Common/Vernacular
Family Scientific Name Names Edible Part Use Reference
Orchidaceae Dactylorhiza romana Roman Dactylorhiza As above Sezik (1967),
(Sebast.) Soó Sezik and Özer
(1983) cited by
Tekinşen and
Güner (2010),
and Sezik (2002)
Orchidaceae Dactylorhiza romana Georgian Orchid As above Ghorbani et al.
subsp. georgica (Klinge) (2014)
Soó ex Renz &
Taubenheim
Orchidaceae Dactylorhiza saccifera Sack-Carrying As above Kreziou et al.
(Brongn.) Soó Dactylorhiza (2015)
Orchidaceae Dactylorhiza sambucina Elder-Flowered Orchid As above Kreziou et al.
(L.) Soó (2015)
Orchidaceae Dactylorhiza umbrosa Persian Marsh-Orchid As above Ghorbani et al.
(Kar. & Kir.) Wendelbo (2014)
Orchidaceae Dendrobium tarberi Rock Lily, King Orchid
Pseudobulbs eaten Steenbeeke
M.A.Clem. & D.L.Jones baked or raw (2001)
Orchidaceae Dipodium Purple Hyacinth Orchid Fleshy starchy thick Steenbeeke
atropurpureum roots are probably (2001)
D.L.Jones edible
Orchidaceae Dipodium Yellow Hyacinth Orchid Fleshy starchy thick Steenbeeke
hamiltonianum roots are probably (2001)
F.M.Bailey edible
Orchidaceae Dipodium pulchellum Dark Hyacinth Orchid Fleshy starchy thick Steenbeeke
D.L.Jones & M.A.Clem. roots are probably (2001)
edible
Orchidaceae Dipodium punctatum Blotched Hyacinth Fleshy starchy thick Steenbeeke
(Sm.) R.Br. Orchid roots are probably (2001)
edible
Orchidaceae Dipodium roseum Pink Hyacinth Orchid Fleshy starchy thick Steenbeeke
D.L.Jones & roots are probably (2001)
M.A. Clem. edible
Orchidaceae Dipodium variegatum Spotted Hyacinth Orchid Fleshy starchy thick Steenbeeke
M.A.Clem. & D.L.Jones roots are probably (2001)
edible
Orchidaceae Disa engleriana NF Tubers process into a Kasulo et al.
Kraenzl. meatless sausage called (2009)
chikande and Kikande
which is consumed as
relish or as snack
Orchidaceae Disa aequiloba Chikanda Mbozi Tubers edible Nyomoro (2009)
Summerh.
Orchidaceae Disa erubescens Rendle Liseki, Liseku, Makaha Starchy tubers used for Mapunda (2007),
Ga Mlutu, Masekele, making salep flour, Challe and Price
Masekeni, Masekendi, chikande meatless (2009), Nyomoro
Mbozi sausage (2009), and
Hamisy (2010)
Orchidaceae Disa fragrans Schltr. NF Tubers processed into Hamisy (2010)
meatless sausage
kikande
(continued)
12 Introduction
Table 1 (continued)
Common/Vernacular
Family Scientific Name Names Edible Part Use Reference
Orchidaceae Disa hamatopetala Baur’s Disa, Ntetemera Tubers processed into Hamisy (2010)
Rendle = Disa baurii meatless sausage
Bolus kikande
Orchidaceae Disa ochrostachya NF As above Mapunda (2007)
Reichb.f. and Hamisy
(2010)
Orchidaceae Disa robusta N.E.Br. Chukande Kijike, As above Mapunda (2007),
Likose, Liisek, Manseke, challe and price
Makaha Ga Mlutu, (2009), Kasulo
Masekele, Masekeni et al. (2009), and
Makaha Ga Mlutu, Hamisy (2010)
Masekele, Masekeni
Makaha Ga Mlutu,
Masekele, Masekeni
Orchidaceae Disa tanganyikensis Chikanda Makete Tubers edible Nyomora (2009)
Summerh.
Orchidaceae Disa ukingensis Schltr. NF Tubers processed into Hamisy (2010)
meatless sausage
kikande
Orchidaceae Disa walleri Rchb.f. Chikanda Mbeya, Tubers edible Nyomora (2009)
Masekelesekele Njombe
Orchidaceae Disa zombica N.E.Br. NF Tubers processed into Kasulo et al.
meatless sausage (2009) and
kikande Hamisy (2010)
Orchidaceae Diuris abbreviata Lemon Doubletail Fleshy starchy tubers Steenbeeke
F. Muell. ex Benth. edible (2001)
Orchidaceae Diuris alba R.Br. White Donkey Orchid Fleshy starchy tubers Steenbeeke
edible (2001)
Orchidaceae Diuris chrysantha Yellow Donkey Orchid Fleshy starchy tubers Steenbeeke
D.L.Jones & M.A.Clem. edible (2001)
Orchidaceae Diuris dendrobioides Purple Donkey Orchid Fleshy starchy tubers Steenbeeke
Fitzg. edible (2001)
Orchidaceae Diuris goonooensis Western Donkey orchid Fleshy starchy tubers Steenbeeke
Rupp edible (2001)
Orchidaceae Diuris lanceolata Lindl. Golden Moths, Snake Fleshy starchy tubers Steenbeeke
Orchid edible (2001)
Orchidaceae Diuris pedunculata Golden Moths, Small Fleshy starchy tubers Steenbeeke
R.Br. Snake Orchid edible (2001)
Orchidaceae Diuris punctata Sm. Purple Donkey Orchid Fleshy starchy tubers Steenbeeke
edible (2001)
Orchidaceae Diuris semilunata Donkey Orchid, Late Starchy tubers edible Cribb and Cribb
Messmer Leopard Orchid, Spotted (1987), Low
Doubletail Orchid (1989), and Low
(1991)
Orchidaceae Diuris striata Rupp NF Fleshy starchy tubers Steenbeeke
edible (2001)
(continued)
Introduction 13
Table 1 (continued)
Common/Vernacular
Family Scientific Name Names Edible Part Use Reference
Orchidaceae Diuris sulphurea R.Br Hornet Orchid, Tiger Starchy tubers edible Anonymous
Orchid, Yellow Tiger (2010a, b),
Orchid Anonymous
(2011), Cribb and
Cribb (1987),
Low (1989), Low
(1991), and
Steenbeeke
(2001)
Orchidaceae Diuris tricolor Fitzg. Tricolor Donkey Orchid Fleshy starchy tubers Steenbeeke
edible (2001)
Orchidaceae Diuris venosa Rupp Veined Doubletail, Fleshy starchy tubers Steenbeeke
Veined Donkey Orchid, edible (2001)
Goat Orchid
Orchidaceae Epipactis royleana Chhasakrungai Boiled roots eaten Sotirov (2015)
Lindl.
Orchidaceae Eriochilus cucullatus Smooth-Leaf Parson’s Tubers edible Steenbeeke
(Labill.) Rchb.f. Bands, Large (2001)
Parson’s-Bands
Orchidaceae Eulophia schweinfurthii Ndulamo, Lisesa Starchy tubers used for Mapunda (2007),
Kraenzl. making salep flour, Challe and Price
chikanda meatless (2009) and
sausage Hamisy (2010)
Orchidaceae Gastrodia procera Large Potato Orchid, Tubers edible Steenbeeke
G.W. Carr Large Cinnamon Bells (2001)
Orchidaceae Gastrodia sesamoides Potato Orchid, Tubers edible Steenbeeke
R.Br. Cinnamon Bells (2001)
Orchidaceae Genoplesium archeri Vriable Midge Orchid Tubers edible Steenbeeke
(2001)
Orchidaceae Genoplesium filiforme NF Tubers edible Steenbeeke
(Fitzg.) D.L.Jones & (2001)
M.A.Clem.
Orchidaceae Genoplesium fimbriata Fringed Midge Orchid Tubers edible Steenbeeke
(R.Br.) D.L.Jones & (2001)
M.A.Clem.
Orchidaceae Genoplesium Dense Midge Orchid Tubers edible Steenbeeke
nudiscapum (Hook.f.) (2001)
D.L.Jones & M.A.Clem.
Orchidaceae Genoplesium nudum Tiny Midge Orchid Tubers edible Steenbeeke
(Hook.f.) D.L.Jones & (2001)
Orchidaceae Genoplesium pedersonii NF Tubers edible Steenbeeke
D.L.Jones (2001)
Orchidaceae Genoplesium rufum Red Midge Orchid Tubers edible Steenbeeke
(R.Br.) D.L.Jones & (2001)
M.A.Clem.
Orchidaceae Glossodia major R.Br. Large Waxlip Orchid Tubers edible Steenbeeke
(2001)
Orchidaceae Habenaria adolphii Chinkanda, Tubers edible Nyomora (2009)
Schultr. Vinying’inya, Songea
(continued)
14 Introduction
Table 1 (continued)
Common/Vernacular
Family Scientific Name Names Edible Part Use Reference
Orchidaceae Habenaria clavata Copper Plant Tubers process into a Kasulo et al.
(Lindl.) Rchb. f. meatless sausage called (2009)
chikande and Kikande
which is consumed as
relish or as snack
Orchidaceae Harbenaria cornuta Chikanda Tubers edible Nyomora (2009)
Lindl.
Orchidaceae Habenaria humilior Chikanda Tubers edible Nyomora (2009)
Rchb.f.
Orchidaceae Habenaria intermedia Intermediate Habenaria Boiled roots eaten Sotirov (2015)
D. Don
Orchidaceae Habenaria keayi NF Tubers and roots are Menzepoh (2011)
Summerh. used to prepare a food
called “napssié” or
ground meat by the
Bagam tribe in the
subdivision of Galim,
Western region of
Cameroon
Orchidaceae Habenaria praestans Chikanda Mbeya Tubers edible Nyomora (2009)
Rendle
Orchidaceae Habenaria xanthochlora Mamkumungu, Starchy tubers used for Challe and Price
Schltr. Manseke, making chikanda (2009) and
Mansekemakubwa, meatless sausage Hamisy (2010)
Mviringo, Likose, Liseke
Orchidaceae Habenaria zambesina NF As above Menzepoh (2011)
Rchb.f.
Orchidaceae Himantoglossum affine NF Starchy tubers used for Sezik (1967),
(Boiss.) Schltr. making salep flour used Sezik and Özer
in ice-cream (1983) cited by
production, Tekinşen and
confectionery and Güner (2010) and
nutritious beverages Ghorbani et al.
(2014)
Orchidaceae Himantoglossum Komper’s Orchid As above Ghorbani et al.
comperianum (Steven) (2014)
Delforge
Orchidaceae Malaxix cylindrostachy Cylindric Raceme Boiled roots eaten Sotirov (2015)
(Lindl.) Kuntze Malaxis
Orchidaceae Neotinea maculata Dense-Flowered Orchid Starchy tubers used for Sezik (2002)
(Desf.) Stearn making salep flour used
in ice-cream
production,
confectionery and
nutritious beverages
Orchidaceae Ophrys climacis NF Starchy tubers used for Deniz (2013)
Heimeier & Perschke making salep flour used
in ice-cream
production,
confectionery and
nutritious beverages
(continued)
Introduction 15
Table 1 (continued)
Common/Vernacular
Family Scientific Name Names Edible Part Use Reference
Orchidaceae Ophrys fusca Link Sombre Bee-Orchid, As above Sezik (1967) and
Dark Bee-Orchid Sezik and Özer
(1983) cited by
Tekinşen and
Güner (2010)
Orchidaceae Ophrys holosericea Late Spider Orchid As above Sezik (1967) and
(Burm. f.) Greuter Sezik and Özer
(1983) cited by
Tekinşen and
Güner (2010)
Orchidaceae Ophrys isaura Renz & NF As above Deniz (2013)
Taubenheim
Orchidaceae Ophrys lycia Renz & Bee Orchid As above Deniz (2013)
Taubenheim
Orchidaceae Ophrys mammosa Desf. Early Spider Orchid As above Tekinşen and
Güner (2010)
Orchidaceae Ophrys phaseliana Sawfly Orchid As above Deniz (2013)
D.Rückbr. & U.Rückbr
Orchidaceae Ophrys scolopax Cav. Woodcock Bee-Orchid, As above Ghorbani et al.
Woodcock Orchid (2014)
Orchidaceae Ophrys sphegodes Mill. Early Spider Orchid As above Ghorbani et al.
(2014)
Orchidaceae Ophrys sphegodes NF As above Ghorbani et al.
subsp. mammosa (Desf.) (2014)
Soó ex E.Nelson
Orchidaceae Orchis adenocheila NF As above Ghorbani et al.
Czerniak (2014)
Orchidaceae Orchis anatolica Boiss. Anatolian Orchid As above Tekinşen and
Güner (2010) and
Citil and Tekinşen
(2011)
Orchidaceae Orchis anthropophora Man Orchid As above Kreziou et al.
(L.) All. (2015)
Orchidaceae Orchis conopea Fragrant Orchid As above Grieve (1971)
Gras = Gymnadenia
conopsea (L.) R.Br.
Orchidaceae Orchis coriophora L. = Fragrant Orchid As above Grieve (1971),
Anacamptis coriophora Tekinşen and
(L.) R.M. Bateman, Güner (2010),
Pridgeon & MW Chase and Citil and
Tekinşen (2011)
Orchidaceae Orchis italica Poir Naked Man Orchid, As above Tekinşen and
Italian Orchid Güner (2010),
Citil and Tekinşen
(2011), and
Kreziou et al.
(2015)
(continued)
16 Introduction
Table 1 (continued)
Common/Vernacular
Family Scientific Name Names Edible Part Use Reference
Orchidaceae Orchis latifolia L. = Early Marsh Orchid As above Grieve (1971)
Dactylorhiza incarnata
(L.) Soó
Orchidaceae Orchis longicruris Naked Man Orchid As above Grieve (1971)
Link = Orchis italica
Poir.
Orchidaceae Orchis maculata L. = Heath Spotted Orchid As above Grieve (1971)
Dactylorhiza maculata
(L.) Soó
Orchidaceae Orchis mascula (L.)L. Early Purple Orchid Starchy tubers used for Ghorbani et al.
salep production (2014)
Orchidaceae Orchis mascula subsp. Early Purple Orchid As above Sezik (1967),
pinetorum (Boiss. & Sezik and Özer
Kotschy) (1983) cited by
E.G.Camus = Orchis Tekinşen and
mascula subsp. mascula Güner (2010),
Grieve (1971),
Hawkes (1944),
and Kreziou et al.
(2015)
Orchidaceae Orchis militaris L. Military Orchid As above Hawkes (1944),
Grieve (1971),
and Kreziou et al.
(2015)
Orchidaceae Orchis morio Green-Winged Orchid As above Hawkes 1944,
L = Anacamptis morio Green-Veined Orchid Grieve 1971, and
(L.) R.M. Bateman, Tekinşen and
Pridgeon & MW Chase Güner (2010),
Citil and Tekinşen
(2011)
Orchidaceae Orchis palustris Jacq. = Toothed Orchid, As above Tekinşen and
Anacamptis palustris Three-Toothed Orchid Güner (2010) and
(Jacq.) R.M. Bateman, Citil and Tekinşen
Pridgeon &MW Chase (2011)
Orchidaceae Orchis provincialis Provence Orchid, Orchis As above Kreziou et al.
Balbis ex Lamarck & De Provence (2015)
DC.
Orchidaceae Orchis pyramidalis L. = Pyramidal Orchid As above Grieve (1971)
Anacamptis pyramidalis
(L.)Rich.
Orchidaceae Orchis saccifera Brong. NF As above Grieve (1971)
= Dactylorhiza
saccifera (Brongn.) Soó.
Orchidaceae Orchis simia Lam Monkey Orchid As above Tekinşen and
Güner (2010) and
Ghorbani et al.
(2014)
Orchidaceae Orchis spitzelii Saut. ex L’orchis De Spitzel As above Sezik (1967),
W.D.J.Koch Sezik and Özer
(1983) cited by
Tekinşen and
Güner (2010)
(continued)
Introduction 17
Table 1 (continued)
Common/Vernacular
Family Scientific Name Names Edible Part Use Reference
Orchidaceae Orchis tridentata Scop. Three-Toothed Orchid Starchy tubers used for Tekinşen and
= Neotinea tridentata making salep flour used Güner (2010) and
(Scop.) R.M. Bateman, in ice-cream Citil and Tekinşen
Pridgeon MW Chase production, (2011)
confectionery and
nutritious beverages
Orchidaceae Orchis ustulata L. = Burnt-Tip Orchid As above Grieve (1971)
Neotinea ustulata (L.)
R.M.Bateman, Pridgeon
& M.W.Chase.
Orchidaceae Orthoceras strictum Horned Orchid, Birds’s Tuberoids probably Steenbeeke
R.Br. Mouth Orchid edible (2001)
Orchidaceae Peristylus constrictus Constricted Peristylus Tubers eaten Sotirov (2015)
Lindley) Lindley
Orchidaceae Platanthera calvigera Club Carrying Boiled roots eaten Sotirov (2015)
Lindl Platanthera
Orchidaceae Platanthera bifolia (L.)Lesser Butterefly Orchid Starchy tubers used for Ghorbani et al.
L.C.Rich making salep (2014)
production
Orchidaceae Prasophyllum brevilabre Short-Lipped Leek Tubers edible Steenbeeke
(Lindl.) Hook.f. Orchid (2001)
Orchidaceae Prasophyllum campestre Starry Leek Orchid Tubers edible Steenbeeke
R.J.Bates & D.L.Jones (2001)
Orchidaceae Prasophyllum Ben Lomond Leek Tubers edible Steenbeeke
caudiculum D.L. Jones Orchid (2001)
Orchidaceae Prasophyllum dossenum NF Tubers edible Steenbeeke
J.Bates & D.L.Jones (2001)
Orchidaceae Prasophyllum elatum Tall Leek Orchid, Piano Tubers edible Steenbeeke
R.Br. Orchid (2001)
Orchidaceae Prasophyllum flavum Yellow leek Orchid Tubers edible Steenbeeke
R.Br. (2001)
Orchidaceae Prasophyllum odoratum Scented Leek Orchid, Tubers edible Steenbeeke
R.S.Rogers Sweet Leek Orchid (2001)
Orchidaceae Prasophyllum patens Broad-Lipped Leek Tubers edible Steenbeeke
R.Br. Orchid (2001)
Orchidaceae Prasophyllum rogersii Marsh Leek Orchid Tubers edible Steenbeeke
Rupp (2001)
Orchidaceae Prasophyllum solstitium Summer Leek Orchid Tubers edible Steenbeeke
R.J.Bates & D.L.Jones (2001)
Orchidaceae Prasophyllum species A Tablelands Leek Orchid Tubers edible Steenbeeke
(2001)
Orchidaceae Pteroglossaspis Mbozi As above Nyomora (2009)
eustachya Rchb.f.
Orchidaceae Pterostylis abrupta Abrupt Greenhood, Tubers edible Steenbeeke
D.L.Jones Drooping Greenhood (2001)
Orchidaceae Pterostylis alata NF Tubers edible Steenbeeke 2001
(Labill.) Rchb.f.
Orchidaceae Pterostylis bicolor Two-Colour Greenhood Tubers edible Steenbeeke 2001
M.A.Clem. & D.L.Jones
(continued)
18 Introduction
Table 1 (continued)
Common/Vernacular
Family Scientific Name Names Edible Part Use Reference
Orchidaceae Pterostylis boormanii Boorman’s Greenhood Tubers edible Steenbeeke 2001
M.A.Clem. & D.L.Jones
Rupp
Orchidaceae Pterostylis chaetophora Hair-Lip Ruddyhood Tubers edible Steenbeeke 2001
M.A.Clem. & D.L.Jones
Orchidaceae Pterostylis coccina Alpen Greenhood Tubers edible Steenbeeke
Fitzg. (2001)
Orchidaceae Pterostylis curta R. Br Blunt Greenhood Tubers edible Anonymous
(2010a, b) and
Steenbeeke
(2001)
Orchidaceae Pterostylis cycnocephala Swan Greenhood Tubers edible Steenbeeke
Fitzg. (2001)
Orchidaceae Pterostylis daintreana NF Tubers edible Steenbeeke
F.Muell. ex Benth. (2001)
Orchidaceae Pterostylis decurva Summer Greenhood Tubers edible Steenbeeke
R.S. Rogers (2001)
Orchidaceae Pterostylis fischii NF Tubers edible Steenbeeke
Nicholls (2001)
Orchidaceae Pterostylis hamata NF Tubers edible Steenbeeke
Blackmore & Clemesha (2001)
Orchidaceae Pterostylis hildae Rainforest Greenhood Tubers edible Steenbeeke
Nicholls (2001)
Orchidaceae Pterostylis laxa NF Tubers edible Steenbeeke
Blackmore (2001)
Orchidaceae Pterostylis longicurva NF Tubers edible Steenbeeke
Rupp (2001)
Orchidaceae Pterostylis longifolia Tall Greenhood Tubers edible Steenbeeke
R.Br. (2001)
Orchidaceae Pterostylis longipetala NF Tubers edible Steenbeeke
Rupp (2001)
Orchidaceae Pterostylis mutica R.Br. Midget Greenhood Tubers edible Steenbeeke
(2001)
Orchidaceae Pterostylis nutans R.Br. Nodding Greenhood Tubers edible Steenbeeke
(2001)
Orchidaceae Pterostylis obtusa R.Br. Jug-Lip Greenhood Tubers edible Steenbeeke
(2001)
Orchidaceae Pterostylis parviflora Tiny Greenhood, Jug Tubers edible Steenbeeke
R.Br. Orchid, Green Snail (2001)
Orchid
Orchidaceae Pterostylis pedunculata Maroonhood Tubers edible Steenbeeke
R.Br. (2001)
Orchidaceae Pterostylis praetermissa NF Tubers edible Steenbeeke
M.A.Clem. & D.L.Jones (2001)
Orchidaceae Pterostylis reflexa R.Br. NF Tubers edible Steenbeeke
(2001)
Orchidaceae Pterostylis setifera NF Tubers edible Steenbeeke
M.A. Clem. (2001)
Orchidaceae Pterostylis species B NF Tubers edible Steenbeeke
(2001)
(continued)
Introduction 19
Table 1 (continued)
Common/Vernacular
Family Scientific Name Names Edible Part Use Reference
Orchidaceae Pterostylis species C NF Tubers edible Steenbeeke
(2001)
Orchidaceae Pterostylis species D NF Tubers edible Steenbeeke
(2001)
Orchidaceae Pterostylis truncate Little Dumpies, Sasauge Tubers edible Steenbeeke
Fitzg. Greenhood (2001)
Orchidaceae Pterostylis woollsii Long-Tailed Greenhood, Tubers edible Steenbeeke
Fitzg. Chinaman Greenhood (2001)
Orchidaceae Roeperocharis Kaloba, Masekel Tubers process into a Challe and Price
wentzeliana Kraenzl meatless sausage called (2009) and
chikande Hamisy (2010)
Orchidaceae Satyrium acutirostrum Chikande As above Hamisy (2010)
Summerh.
Orchidaceae Satyrium amblyosaccos NF Tubers process into a Kasulo et al.
Schltr. meatless sausage called (2009)
chikande and kikande,
which is consumed as
relish or as snack
Orchidaceae Satyrium atherstonei Chikande Ligos, Ingingi, Starchy tubers used for Mapunda (2007),
Rchb.f. = Satyrium Jike, Lidala, Lisekejike, making salep flour, Challe and Price
trinerve Lindl. Lisekeni, Lisekenilidala, chikanda meatless (2009) and
Madala, sausage Hamisy (2010)
Masekenimadala,
Numbunumbu, Sidala,
Visekenividala, Vijike
Orchidaceae Satyrium breve Rolfe Chikande Tubers edible Nyomora (2009)
Orchidaceae Satyrium buchananii Dochamua, Ligosi, Tubers process into a Mapunda (2007),
Schltr. Likosi, Lisekedochamua, meatless sausage called Kasulo et al.
Lisekedume, chikande and (2009) and
Lisekekiume, Magosi, kikande,which is Hamisy (2010)
Masekenidume, consumed as relish or
Masekeni magosi, as snack, starchy tubers
Sisekeni sigosi, used for making salep
Titisigosi, flour
Visekenivikhosi,
Visekenivigosi
Orchidaceae Satyrium carsonii Rolfe NF Tubers process into a Kasulo et al.
meatless sausage called (2009)
chikande and kikande,
which is consumed as
relish or as snack
Orchidaceae Satyrium chlorocorys Chikanda Jike Rukwa Tubers edible Nyomora (2009)
Rolfe
Orchidaceae Satyrium crassicaule Chikanda, Kikande, Tubers process into a Nyomora (2009)
Rendle Makete, Simbegi meatless sausage called and Hamisy
chikande and kikande, (2010)
which is consumed as
relish or as snack
Orchidaceae Satyrium nepalense Nepal Satyrium Pseudobulbs eaten Sotirov (2015)
D.Don boiled
Orchidaceae Satyrium robustum Kidume Tubers processed into Hamisy (2010)
Schltr. meatless sausage
kikande
(continued)
20 Introduction
Table 1 (continued)
Common/Vernacular
Family Scientific Name Names Edible Part Use Reference
Orchidaceae Satyrium sacculatum Chikanda, Kikande, Tubers edible Nyomora (2009)
(Rendle) Rolfe Masehelesehele, Vikan,
Dokando, Songea
Orchidaceae Satyrium volkensii Volkens ‘Satyrium Tubers processed into Hamisy (2010)
Schltr. meatless sausage
kikande
Orchidaceae Serapias vomeracea ssp. NF Starchy tubers used for Tekinşen and
orientalis making salep Güner (2010),
Grueter = Serapias Citil and Tekinşen
orientalis (Greuter) (2011), Sezik
H. Baumann & Künkele (1967), Sezik and
Özer (1983) cited
by Tekinşen and
Güner (2010)
Orchidaceae Steveniella satyrioides NF Starchy tubers used for Ghorbani et al.
(Spreng.) Schltr making salep (2014)
Orchidaceae Thelymitra carnea R.Br. Tiny Sun Orchid, Pink Tubers edible Steenbeeke
Sun Orchid (2001)
Orchidaceae Thelymitra circumsepta Naked Sun Orchid Tubers edible Steenbeeke
Fitzg. (2001)
Orchidaceae Thelymitra cyanea Veined Sun Orchid Tubers edible Steenbeeke
(Lindl.) Benth. (2001)
Orchidaceae Thelymitra fragrans NF Tubers edible Steenbeeke
D.L.Jones & M.A.Clem. (2001)
Orchidaceae Thelymitra Scented Sun Orchid Tubers edible Steenbeeke
megacalyptra Fitzg. (2001)
Orchidaceae Thelymitra nuda R.Br. Plain Sun Orchid Tubers edible Steenbeeke
(2001)
Orchidaceae Thelymitra pauciflora Slender sun Orchid Tubers edible Steenbeeke
R.Br. (2001)
Orchidaceae Tipularia discolour Crane Fly Orchid, Starchy corm eaten Sotirov (2015)
(Pursh) Nuttall Crippled Cranefly Orchid
Oxalidaceae Oxalis corymbosa DC. = Pink Shamrock Sweet crisp swollen tap Cribb and Cribb
Oxalis debilis var. root eaten (1987)
corymbosa (DC.)
Lourteig.
Oxalidaceae Oxalis deppei Lodd. ex Orach, Deppei Wood Fleshy root eaten Fernald et al.
Sweet Sorrel boiled in Mexico (1985) and
Facciola (1990)
Oxalidaceae Oxalis perennans Haw. Grassland Wood Sorrel Small tap roots eaten Low (1991)
by aborigines
Oxalidaceae Oxalis radicosa A.Rich. Dwarf Wood Sorrel Small tap roots eaten Low (1991)
= Oxalis corniculata L. by aborigines
Oxalidaceae Oxalis sp. Wood Sorrel Small tap roots eaten Low (1991)
by aborigines
Oxalidaceae Oxalis stricta L. Yellow Wood Sorrel Roots edible Yanovsky (1936),
Uphof (1968),
and Facciola
(1990)
(continued)
Introduction 21
Table 1 (continued)
Common/Vernacular
Family Scientific Name Names Edible Part Use Reference
Oxalidaceae Oxalis tuberosa Molina Oca Tubers eaten boiled, Facciola (1990),
roasted or candied, one Groen et al.
of the principal food (1996), Flores
crop of the Indians in et al. (2003), van
the Andes, second to Wyk (2006), and
potato, tubers can be Santich et al.
eaten unpeeled, raw, in (2008),
salads, pickled, boiled,
fried or in soups and
stews. Also can be
dried in sun to sweeten
up
Oxalidaceae Oxalis violacea L. = Violet Wood Sorrel Roots edible Yanovsky (1936)
Oxalis debilis var. and Facciola
corymbosa (DC.) (1990)
Lourteig.
Piperaceae Piper methysticum L.f. Kava, Kava Kava, Awa, The root and rhizome McDonald and
Kava Pepper (underground stem) of Jowitt (2000) and
kava are used to NCAM (2006)
prepare beverages,
extracts, capsules,
tablets and topical
solutions. Kava is
widely and commonly
consumed as a social
beverage to establish
kinship in the Pacific
island communities
Poaceae Agropyron repens (L.) English Couch, Quick Roots ground into a Cribb and Cribb
P. Beauv. = Elymus Grass meal and used to make (1987)
repens (L.) Gould bread
Poaceae Bambusa pallida Munro Mokal Bah (Assamese); Young rhizomes are Patiri and Borah
Mai Phiu, Phai Song eaten as vegetable after (2007)
Kham (Thai) processing
Poaceae Bambusa tulda Roxb. Spineless Indian As above Patiri and Borah
Bamboo, Calcutta Cane; (2007)
Jati Bah (Assamese)
Poaceae Chrysopogon Vetiver Grass, Khas- Vetiver is used Balasankar et al.
zizanioides (L.) Roberty Kahs, Khus Grass domestically in (2013)
cooking; it’s infused in
tea and also used in
baking
Poaceae Eleusine indica (L.) Indian Goosegrass, Roots edible raw Kunkel (1984)
Gaertn. Wiregrass,
Crowfootgrass
Poaceae Hordeum bulbosum L. Abu Suwaif Bulbous roots chewed Tanaka (1976)
and eaten occasionally and Facciola
(1990)
(continued)
22 Introduction
Table 1 (continued)
Common/Vernacular
Family Scientific Name Names Edible Part Use Reference
Poaceae Imperata cylindrica L. Blady Grass, Alang- Aborigine children Burkill (1966),
Alang, Lalang Wooly sucked the roots, Cribb and Cribb
Grass underground shoots (1987), Low
like sugarcane, used as (1991), and Hu
survival food, a kind of (2005)
beer is made from the
roots in Peninsular
Malaysia China: fresh
rhizome chewed by
rural people for the
sweet juice
Poaceae Phragmites australis Common Reed Grass Rhizomes coked like Tanaka (1976),
(Cav.) Trin. Ex Steud. potatoes Fernald et al.
(1985), and
Facciola (1990)
Poaceae Saccharum spontaneum Wild Cane, Wild Young shoots and Patiri and Borah
L. Sugarcane, Fodder Cane; rhizomes are eaten as (2007)
Khagori (Assamese); sugarcane, sweet in
Mojora (Mishing) taste
Poaceae Triticum repens L. = Couch Grass, Twitch, In Norway, roots Sayce (1953)
Elymus repens (L.) Quick Grass, Quitch ground into flour
Gould. Grass, Dog Grass
Poaceae Vetiveria nigritana Black Vetiver grass; Nigeria (Kano State, Mortimore (1989)
(Benth.) Jema (Huasa) northern): roots eaten.
Stapf = Chrysopogon
nigritanus (Benth.)
Veldkamp.
Poaceae Vetiveria zizanioides Vetiver Grass, Khas- Roots used to prepare Pareek and
(L.) Nash = Chrysopogon Khas Grass, Akar Wangi sherbets or soft drinks Kumar (2013)
zizanioides (L.) Roberty. (Malay) during summer in and Balasankar
Nothern India. Vetiver et al. (2013)
oil is used for
flavouring syrups,
ice-cream and food
preservation. Khus
essence is used in cool
drinks, for reducing
pungency of chewing
tobacco preparations
and to provide sweet
note to other
masticatories
Vetiver roots is used
domestically in
cooking; it’s infused in
tea and also used in
baking
Poaceae Zizaniopsis miliacea Water Millet, Southern Young rhizome, cut Gibbons and
(Michx.) Döll & Asch. Wild Rice into pieces cooked and Tucker (1979),
served with butter Fernald et al.
(1985), and
Facciola (1990)
(continued)
Introduction 23
Table 1 (continued)
Common/Vernacular
Family Scientific Name Names Edible Part Use Reference
Rubiaceae Ixora subsessilis Wall. Dieng Jowat Flower, shoot, root Sawian et al.
ex G.Don (Meghalaya); Nang Guo eaten in Meghalaya (2007)
Long Chuan Hua
(Chinese)
Rubiaceae Morinda officinalis F.C. Chinese Herbal Morinda, In China, fresh or dried Hu (2005), Li et
How Medicinal Indian roots are cooked with al. (2009)
Mulberry, Morinda Root pork for a broth as
health food called
Chinese bupin (Hu
2005). In South China,
Hong Kong and Macao,
this plant has been
developed into various
health foods, such as
‘Ba-ji-tian wine’,
‘Ba-ji-zi-bu Gao’ (Li
et al. 2009)
Rubiaceae Paederia foetida L. Rekang Nemthu Roots eaten in Karbi Kar and
Assam Borthakur (2008)
Rubiaceae Paederia stenobotrya White Paedaria Root ground with Hu (2005)
Merr. soaked soybean for a
milk, cooked and given
to people with jaundice
Simaroubaceae Eurycoma longifolia Tongkat Ali (Malay) Traditional users of Low et al. (2013)
Jack Tongkat Ali brew tea
from the dried chips of
the tree’s root. Tea,
coffee and carbonated
beverages, pre-mixed
with the root extract are
available commercially
for the improvement of
general health and
libido in Malaysia
NF not found
is processed into nutritious and wholesome from the genera Disa, Habenaria and Satyrium.
beverages, confectionary and ice-cream. Salep is Many species in the orchid genera Pterostylis,
especially popular in Middle East and some parts Diuris and Thelymitra have edible tubers. Other
of Europe. The major salep producing orchid orchid species with edible roots and tubers are
species are from the genera Anacamptis, Orchis, listed in Table 1.
Ophrys, Disa, Dactylorhiza and Himantoglossum.
In Africa, edible orchid tubers are collected from
the wild and processed into a meatless sausage References
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Iris x germanica
Chinese: Déguó Yuānwěi tile hybrid between I. pallida and I. variegata L.,
Czech: Kosatec Německý both of which also have the chromosome number
Danish: Have-Iris, Iris, Sværdlilie 2n = 24 (Henderson 1992, 2002).
Dutch: Blauwe Lis, Duitse Lis, Duitse Lis Sort,
Lis Sort
Esperanto: Irido Ĝardena, Irido Germana Agroecology
Estonian: Aediiris
Finnish: Saksankurjenmiekka, Sininen Iris x germanica thrives in full sun, in fertile,
Kurjenmiekka well-drained, neutral to slightly acidic soil with
French: Flambe, Iris, Iris Allemande, Iris pH 6–7. It needs light watering. It is widely
d’Allemagne, Iris Germanique grown as a garden ornamental in the temperate
German: Deutsche Schwertlilie, Echte regions of Australia and has become naturalised
Schwertlilie, Gelbe Schwertlilie, along roadsides, in waste areas and in bushland.
Himmelschwertel, Ritter-Schwertlilie, Türk
Schwertlilie
Hungarian: Kerti Nőszirom, Kék Nőszirom, Edible Plant Parts and Uses
Nepáli Nőszirom
India: Puskaramulam (Malayalam), Kombirei Orrisroot is often included as one of the many
(Manipuri), Haimavati, Mulam, Padma- ingredients of Ras el hanout, a blend of herbs
Pushkara, Parasikavaca, Puskaramulam and spices used across the Middle East and
(Sanskrit) North Africa, primarily associated with
Italian: Giaggiolo Paonazzo, Giaggiolo Moroccan cuisine (Surhone et al. 2011). Peeled
Maggiore rhizomes of I. germanica (orrisroots) can be
Japanese: Ayame, Hanashoubu, Kakitsubata used as a flavouring in ice cream, confectionery
Korean: Ailiseu Germanica and baked goods (Bender 2009). Orris is also an
Persian: Bikh-I-Banafshah ingredient in many brands of gin. Orrisroot has
Polish: Kosaciec Bródkowy, Kosaciec Niemiecki, been used in tinctures to flavour syrups; its taste
Kosaciec Ogrodowy is said to be indistinguishable from raspberry
Portuguese: Lírio-Cardano, Lírio-Cardeno, (Chase 1900). In France, starch of I. germanica
Lírio-Da-Alemanha, Lírio-Germânico, root had been recommended a famine food for
Lírio-Roxo extending bread flour, after removal of the bitter
Slovašcina: Bradata Perunika, Nemška Perunika, element (Parmentier 1781, cited by Freedman
Perunika Nemška 2009).
Slovencina: Kosatec Nemecký
Spanish: Iris, Lirio Cardeno, Lirio Común
Swedish: Trädgårdsiris Botany
Thai: M̀āntā Germanica
Tibetan: Su Dag Dkar Po Iris x germanica is a rhizomatous, perennial herb,
Turkish: Mor Süsen, Navruzu, Türk Süzeni growing to 120 cm high, forming a large clump
Vietnamese: Diên Vỹ German hoa tím to 30 cm wide (Plate 1). Rhizomes are homoge-
Welsh: Gellesgen Farfog neous, creeping on soil surface or to 10 cm depth,
usually many-branched, light brown, 1.2–2 cm in
diameter and smooth, with nodal rings; branches
may arise in the fan or as many as 15–20 nodes
Origin/Distribution are produced prior to active leaves. Stems are
green, 2–3-branched, solid, 60–120 cm × 1–1.5 cm
I. x germanica probably originated in central and glaucous. Leaves are purplish at base and
southern Europe and the Balkan Peninsula. Iris folded midrib to base, glaucous, ensiform and
germanica is considered to be a natural and fer- 45 cm long by 3.5 cm wide. Inflorescences with
Iris x germanica 29
Nutritive/Medicinal Properties
Rhizome Phytochemicals
along with a known compound, nigricin 4′-(β-D- of isoflavone glucosides. Two novel compounds
glucopyranoside), were isolated from Iris ger- 5,2-dihydroxy-3-methoxy-6,7-
manica rhizomes (Atta-ur-Rahman et al. 2003a). methylenedioxyflavone and 5,7,2-trihydroxy-
Nine isoflavonoids: 5,7-dihydroxy-3-(3′-hydroxy 6-methoxyflavanone were isolated from the
-4′,5′dimethoxy)-8-methoxy-4H-1-benzopyran- fraction B of the rhizome chloroform extract
4-one; 5,7-dihydroxy-3-(3′-hydroxyl-4′, 5′-dim- (Singab et al. 2006). The methanol extract of Iris
ethoxy)-6-methoxy-4H-1-benzopyran-4-one; 5, germanica rhizome afforded one new compound,
7-dihydroxy-3-(4′-hydroxy)-6-methoxy-4H-1- 6,6-ditetradecyl-6,7-dihydrooxepin-2(3H)-one
benzopyrane-4-one; 5-hydroxy-3-(4′-hydroxy)- and five known compounds, 1-(2-(6′-hydroxy-2′-
6,7-methylenedioxy-4 H -1-benzopyran-4-one; methylcyclohex-1′-enyloxy)-5-methoxyphenyl)
5-hydroxy-3-(4′-methoxy)-6,7-methylenedioxy-4H- ethanone; 4-hydroxy-3-methoxyacetophenone;
1-benzopyran-4-one; 5-methoxy-3-(4′-hydroxy)- irisolone; irisolidone and 2-acetoxy-3,6-
6,7-methyenedioxy-4 H -1-benzopyran- 4-one; dimethoxy-1,4-benzoquinone (Asghar et al.
5,7-dihydroxy-3-(3′-hydroxy-4′-methoxy)-6- 2009).
methoxy-4H-1-benzopyran-4-one; 5,7-dihydroxy A total of 20 compounds identified as isofla-
-3-(3′-methoxy-4′-hydroxy)-6-methoxy-4H-1- vones, isoflavone glycosides and acetovanillone
benzopyran-4-one and isopeonol were isolated were isolated from the lipophilic and polar
from the rhizome (Atta-ur-Rahman et al. 2003b). extracts of Iris germanica rhizomes (Schütz et al.
Five new di- and triglycosides, irigenin 7-[O-β-D- 2011). A new isoflavone glycoside, iriflogenin-
glucopyranosyl-(1 → 6)- β - D - glucopyranoside] 4′-O-gentiobioside, was isolated. Nine isofla-
(1); 6-hydroxygenistein 4′-[O-β-D-glucopyranosyl- vones (totalling 180 mg/g) were identified in I.
(1 → 6)-β-D-glucopyranoside] (2); nigricin germanica resinoid: irigenin, iristectorigenin A,
4′-[O-β-D-glucopyanosyl-(1 → 6)-β-D- nigricin, nigricanin, irisflorentin, iriskumaonin
glucopyranoside] (3); nigricin 4′-[O-β-D- methyl ether, irilone, iriflogenin and irisolidone
glucopyanosyl-(1 → 2)-O-[α-L-rhamnopyranosyl- (Roger et al. 2012). The methanolic extract of
(1 → 6)]-β-D-glucopyranoside] (4) and 7-{4′- Iris germanica rhizomes yielded two new
{[2″-O-(4′′′′-acetyl-2′′′′-methoxyphenyl)-β-D- compounds, irigenin S and iriside A, together
glucopyranosyl]oxy}-3′-( β - D -glucopyran- with known compounds: stigmasterol, α-irone,
osyloxy)phenyl]-9-methoxy-8H-1, γ-irone, 3-hydroxy-5-methoxyacetophenone,
3-dioxolo[4,5-g]-[1 benzopyran-8-one-] (5), irilone, irisolidone, irigenin, stigmasterol-3-
along with a known compound, nigricin 4′-(β-D- O-β-D-glucopyranoside, irilone 4′-O-β-D-
glucopyranoside), were isolated from Iris ger- glucopyranoside and iridin (Ibrahim et al. 2012).
manica rhizomes (Nasim et al. 2003). Irigenin Two new compounds, 5-methoxy-3′,4′-
(5,7,3′-trihydroxy-6,4′,5′-trimethoxyisoflavone) dihydroxy-6,7-methylenedioxy-4H-1-
and iristectorigenin A (5,7,3′-trihydroxy-6,4′- benzopyran-4-one (iriskashmirianin A) and
dimethoxyisoflavone) along with their 7-O-β-D- 5,3′-dihydroxy-3-(4′-β-D-glucopyranosyl)-6,7-
glucosides, iridin and iristectorin A, respectively, methylenedioxy-4H-1-benzopyran-4-one (ger-
were found as the major components in adventi- manaism H), along with eight known compounds,
tious roots in the liquid medium, and the total namely, irilone, iriskumaonin methyl ether, iriflo-
isoflavone content was about 3.6 μmol per g fresh genin, irisolone, irifloside, irilone 4′-O-β-
weight in 3-week-old I. germanica cultures glucopyranoside, germanaism A and germanaism
(Akashi et al. 2005). The total isoflavone content B were isolated from the rhizomes (Xie et al.
(glucosides + aglycones) at 6 h after the start of 2013). New ceramides irisamides A and B and
CuCl2 treatment was 1.4-fold the initial value, isoflavone iridin S were isolated from the rhi-
and nearly the same content was maintained for zome (Mohamed et al. 2013). The methanol
48 h. Thus, the main effect of CuCl2 treatment extract of Iris germanica afforded a new benzene
appeared to be the induction of hydrolysis derivative 2′-methyl-6′-hydroxy cyclohexenyl-
32 Iridaceae
3-methyl-1-acetophenone ether and another (0.14 %); benzoic acid, 4-ethoxy-, ethylester
known benzene derivative isopenol together with (0.12 %); hexadecane (0.11 %); thiosulfuric acid,
two known isoflavones, irisolone and irisolidone S-(2aminoethyl) ester (0.1 %); tetradecane
(Asghar et al. 2010). Twenty-one compounds (0.08 %); and decane (0.06 %).
were isolated from the rhizomes of Iris german- Orrisroot is used as an aromatic agent. Iris
ica and identified as ombuin (1); 5, 3, 3′-trihy- pallida is the best for extractive purposes
droxy-7, 4′-dimethoxyflavanone (2); naringenin followed by I. germanica and I. florentina for
(3); cirsiliol-4′-glucoside (4); 3β, 4′-dihydroxy- botanical sources of orrisroot preparations (con-
7,3′-dimethoxyflavonone-5-O-β-D-- crete, liquid, oil, root extract) (Fenaroli 1994).
glucopyranoside (5); genistein (6); irilin D (7); Orris absolute consists mainly of isomeric irones
muningin (8); 5, 7, 4′-trihydroxy-6, 3′, 5′-trime- (Guenther 1952). Orris absolute (alcoholic
thoxyisoflavone (9); tectorigenin (10); irigenin extraction or distillation of orris concrete) was
(11); tectoridin (12); iridin (13); mangiferin (14); reported to have a yield of 0.03–0.05 % dried rhi-
irisxanthone (15); pyroglutamic acid (16); 2, 4′, zome and to contain 55–58 % ketones (irones),
6-trihydroxy-4-methoxybenzophenone-2-O-β-D- free or partially esterified fatty acids (methyl
glucoside (17); apocynin (18); androsin (19); myristate, methyl caprylate, methyl laurate,
β-sitosterol (20); and daucosterol (21) (Xie et al. methyl oleate, methyl linoleate), aldehydes
2014). (oleic, benzoic), alcohols (benzylic), terpene and
Eleven compounds were identified in the petro- sesquiterpene (Burdock 1994). Orris resinoid
leum ether extract (oil) of I. germanica: 9-hexa- (yield 1–3.3 %) was reported to contain 62–78 %
decanoic acid methyl ester; 9-octadecenoic acid ketones (irones) (Burdock 1994). Orris absolute
methyl ester; 8-octadecenoic acid methyl ester; prepared from Iris x germanica was reported to
11-octadecenoic acid methyl; 10-octadecenoic contain (Z)-a-irone (558.4–62.7 %), (Z)-γ-irone
acid methyl ester; 13-octadecenoic acid methyl (32.2–36.9), (E)-α-irone (1.7–3.9 %) and (Z)-β-
ester; 16-octadecenoic acid methyl ester; 1, irone (0.4–2.3 %) and orris essential oil to con-
2-benzenedicarboxylic acid diisooctyl ester; bis(2- tain (Z)-α-irone (57.6–66.2 %), (Z)-γ-irone
ethylhexyl) phthalate; methyl 6-methyl heptano- (33.8–39.4 %), (E)-α-irone (0.5–2.7 %) and (Z)-
ate; and nonanoic acid, 9-oxo-methyl ester (Asghar β-irone (0–2.2 %) (Galfre et al. 1993). (+)-cis-γ-
et al. 2011). Twenty-three compounds were identi- Irone and (+)-cis-α-irone were found to have the
fied in the essential oil of Syrian Iris germanica most interesting organoleptic properties. Orris
with myristic acid (61.42 %) as the major compo- essential oil is a cream-coloured solid and is
nent (Almaarri et al. 2013). The other sub-major often referred to as orris butter and often and
compounds obtained were elaidic acid methyl erroneously referred to as orris concrete which is
ester (9-octadecenoic acid methyl ester) (6.61 %), a dark, viscous material perhaps more correctly
lauric acid (5.69 %), octadecanoic acid methyl referred to as a resinoid.
ester (stearic acid methyl ester) (5.40 %), palmitic Irones extracted from Iris species are aromatic
acid methyl ester (5.10 %) and palmitic acid principles used in the perfume and flavour indus-
(4.87 %). The minor compounds were linoleic try to impart violet fragrance. Four major irone
acid methyl ester (1.19 %); linoleic acid (1.1 %); compounds (trans α-, cis α-, cis γ- and β-irones)
docosane (1.08 %);10-octadecenoic acid, methyl had been described (Naves et al. 1947; Schinz
ester (0.7 %); 3-methyl-1,2-benzoisoselenazple et al. 1947; Jaenick and Marner 1990) as respon-
(0.61 %); nonadecane (0.55 %); oleic acid sible for the characteristic scent of Iris oil. Natural
(0.53 %); sulphurous acid,2-propyl tetradecyl irones (terpenoids) in essential oil of Iris rhizomes
ester (0.28 %); α-irone (0.25 %); 2-propenoic were found to be formed by oxidative degrada-
acid, 3-4(-methoxy-phenyl)-2ethyl-hexyl ester tion of other higher terpenes C31-triterpenoids
(0.16 %); 1,2-benzenedicarboxylic acid, dioctyl produced by the plant (Krick et al. 1984).
ester (0.15 %); decanoic acid (capric acid) Triterpenoids called iridals were shown to be
Iris x germanica 33
irone precursors (Jaenick and Marner 1990). manica rhizomes (Bonfils and Sauvaire 1996).
Iridals originated from C30 olefin squalene Monocycloiridals were mostly represented in
(Jaenick and Marner 1990, Marner et al. 1988; young rhizomes, whereas cycloiridals appeared
Marner 1997). The incorporation of [2-14C]ace- to be the main iridal formed in the old ones. The
tate, [2-14C]mevalonate and [3H]squalene proved study of the ratio of free iridals (FI)/iridal esters
the squalenoid nature of the iridals (Marner et al. (IE) revealed a higher ratio in the youngest rhi-
1988). Methionine was readily incorporated into zomes. In addition 20 irone-related compounds
cycloiridals of Iris pallida dalmatica, thus indi- containing 10-16C-atoms were detected in orris
cating that the methylation of iridals via concrete of Moroccan origin (I. germanica)
S-adenosyl-L-methionine led to the formation of (Maurer et al. 1989). Iris germanica was found to
the irone moiety of the bicyclic compounds. The have a fresh mass of 1342 g/plant and a dry mass
cycloiridals, C31-triterpenoids, also served as of 437 g/plant and to contain 595 mg/kg DM
precursors of the irones (Marner et al. 1990). irone content that was composed of 79.1 % cis-γ-
Since 1982, more than 25 different iridal struc- irone and 20.9 % cis-α-irone (Firmin et al. 1998).
tures were characterised (Jannick and Marner An epoxidised iridal 18,19-epoxy-10-deoxyiridal
1990, Abe et al. 1991; Marner and Kerp 1992) was isolated from rhizomes of Iris germanica
and separated into 3 classes monocycloiridals, var. ‘Rococo’ (Bonfils et al. 1998). Ten enantio-
bicycloiridals and spiroiridals. Rhizomes of both pure isomers of irone were reported to be the
I. pallida varieties were found to contain 0.1– odoriferous principle of orrisroot oil, by means of
1.1 %, and I. germanica 0.1–0.5 % of iridals and enzyme-mediated approaches, starting from
their esters (Marner and Kerp 1992). Bonfils commercial racemic Irone alpha® (Brenna et al.
et al. (1996) found iridals were membrane 2003). A new iridal irigermanone and nine known
located. Two triterpenoids iripallidal and iriflo- congeners were isolated from the dried rhizomes
rental, known irone precursors, were found to be of German iris (Iris germanica) (Potterat et al.
solubilised by phosphatidylcholine. These cyclo- 2014). The compound was a structurally unique
iridals were found to be integrated within liposo- noriridal and possessed an unprecedented meth-
mal membranes and appeared to have a structural ylcarbonyl group instead of the α,β-unsaturated
role within cells comparable to that of sterols. aldehyde function typical for this group of
Iris germanica had high levels of (+)cis-α-irone triterpenes.
and low content of trans-α-irone, while I. pallida
had a high content of (-) cis-α-irone (Maurer
et al. 1989; Firmin et al. 1998). Dehydration of Other Plant Parts’ Phytochemicals
rhizome slices led to an increase of iridal esters
with a concomitant drop of free iridals, as well as A C-glycosyl flavone, embinin, is isolated from
phospholipids, in the drought-resistant species, the petals of I. germanica (Kawase and Yagishita
Iris germanica (Bonfils et al. 1994). Analysis of 1968). In Iris germanica, flower colour was
the intracellular membranes of Iris germanica found to be determined by two distinct biochemi-
rhizomes indicated high amounts of iridals when cal pathways, the anthocyanin biosynthetic and
compared to sterols and lower sterol amounts the carotenoid biosynthetics (Warburton et al.
than in other plant microsomes (Bonfils et al. 1978; Meckenstock 2005). In cyanic iris flowers,
1995). Incubation of a monocyclic iridal (a triter- the main pigments reported were delphinidin-
penoid from Iris species) with rat liver micro- type anthocyanins imparting blue, violet and
somes in the presence of NADPH led to the maroon colours, while in acyanic flowers, the
degradation of this substrate and to the genera- main pigments were carotenoids, producing
tion of 16-hydroxy-iridal (isoiridogermanal) orange, yellow and pink colours. A red iris does
(Bonfils et al. 1955). Iridal composition was also not naturally exist because of a deficiency in the
found to differ, depending on the age of Iris ger- synthesis of pelargonidin- and cyanidin-type
34 Iridaceae
anthocyanins, usually responsible for red and pounds was rated in the following order:
orange hues or that red carotenoids, such as lyco- BHA > BHT > α-tocopherol > > aqueous iris
pene, do not accumulate in large concentrations, extract > ethanol iris extract. The scavenging
thus producing only pink flowers, as opposed to effect of the aqueous and ethanol iris extracts and
red ones (Jeknić et al. 2014). With a goal of the reference compounds on the DPPH* radical
developing iris cultivars with red flowers, they decreased in the following order:
transformed a pink iris I. germanica, ‘Fire Bride’, BHA > α-tocopherol > BHT > aqueous
with a bacterial phytoene synthase gene (crtB) iris > ethanol iris extract. These results indicated
from Pantoea agglomerans under the control of that both iris extracts produced a noticeable scav-
the promoter region of a gene for capsanthin– enging of free radicals; free radical scavenging
capsorubin synthase from Lilium lancifolium activity also increased with the concentration of
(Llccs). They showed that ectopic expression of a iris extracts. The metal chelating effect of both
crtB could be used to successfully alter the colour iris extracts and the reference compounds
of certain flower parts in I. germanica ‘Fire decreased in the following order: aqueous iris
Bride’ and produce new flower traits. The colour extract > ethanol iris extract > BHA > α-tocophero
changes were confirmed primarily due to the l > BHT. The results indicated that iris had in-
accumulation of lycopene. Other carotenoids and vitro antioxidant properties, which could be the
chlorophyll found in both control and crtB peri- major factor responsible for the inhibition of lipid
gon (standards and falls), ovaries and flower stalk peroxidation. The petroleum ether extract (oil) of
were neoxanthin, violaxanthin, lutein, lycopene, I. germanica exhibited good DPPH scavenging
chlorophyll a, chlorophyll b and α + β-carotene. activity but showed moderate inhibition in lin-
Carotenoids β-carotene 7.1 % (4.3 mg), lutein oleic acid oxidation as compared to BHT (Asghar
87.8 % (53.2 mg), neochrome (zeaxanthin furan- et al. 2011). The radical scavenging activity of
oxide) 3.5 % (2.1 mg) and 7 minor constituents the ethanolic extracts of the aerial parts and rhi-
1.6 % (1 mg) were found in the green aerial parts zomes of I. germanica were found to have IC50
of I. germanica (Bucheckerand and Liaaen- values of 5.38 and 12.3 mg/ml, respectively,
Jensen 1975). From the leaves of I. pallida, 0.1– whereas total antioxidant activity of the extracts
0.7 % of iridals and their esters were extracted (at 3.15 mg/ml) was 98.7 % and 97.4 %, respec-
and from I. germanica leaves 0.05–0.3 % (Marner tively, and total phenolic content was 267.36 and
and Kerp 1992). 331.96 8 mg gallic acid equivalent/g extract,
respectively (Burcu et al. 2014). HPLC analysis
of phenolic compounds identified protocatechuic
Antioxidant Activity acid (0.356 mg/g extract), chlorogenic acid
(0.164 mg/g extract) and ferulic acid (0.164 mg/g
Both aqueous and ethanol I. germanica rhizome extract) as the main phenolic acids contained in
extracts exhibited strong total antioxidant activ- the extract of the aerial parts of I. germanica,
ity, showing 95.9, 88.4, 79.9 % and 90.5, 78.0, whereas chlorogenic acid (2.44 mg/g extract),
65.3 % inhibition on peroxidation of linoleic acid (+)-catechin (2.14 mg/g extract) and ferulic acid
emulsion at concentrations of 10, 30 and 50 μg/ (0.452 8 mg/g extract) were identified as the main
ml, respectively, and in all cases exceeded the phenolic acids contained in the extract of the
effect of 30 μg/ml α-tocopherol solution (36.6 %) rhizome.
(Nadaroğlu et al. 2007). Both extracts also pos-
sessed effective reducing power and exhibited
free radical scavenging, superoxide anion radical Anticancer Activity
scavenging, hydrogen peroxide scavenging and
metal chelating activities at concentrations of 20, Isoflavones from I. germanica rhizomes, irigenin,
40 and 60 μg/ml. The reducing power of aqueous irilone and iriskashmirianin exhibited moderate
and ethanol iris extracts and reference com- activity as inducers of NAD(P)H:quinone reduc-
Iris x germanica 35
tase (QR) in cultured mouse Hepa 1c1c7 cells, (EAC) cancer cell line with IC50 of 20.9 7 and
with CD values (concentration required to double 4.3 μM for 3-(4, 5-dimethylthiazole-2-yl)-2,
the specific activity of QR) of 3.5–16.7 μM, 5-diphenyltetrazoli-umbromide (MTT) and ATP
whereas weak activity was observed with com- assay methods, respectively (Xie et al. 2013).
pounds iriflogenin and iriskashmirianin in the
radical (DPPH) scavenging bioassay (IC50 values
89.6 and 120.3 μM, respectively) (Ito et al. 1995). Antimutagenic Activity
Iridals extracted from Iris germanica exhibited
antitumour effects in-vitro against human tumour The ethanolic extract of the aerial parts of I. ger-
cell lines: A2780 and K562 (and for each one, a manica exhibited antimutagenic effects at 3 and
drug-sensitive and a drug-resistant cell line) with 0.3 mg/plate concentrations as assessed using the
IC50 values of 0.1–5.3 μg/ml (Bonfils et al. 2001). Ames Salmonella typhimurium/microsome
Some of them were shown to be more effective mutagenicity test (Burcu et al. 2014). The rhi-
than doxorubicin. Studies by Halpert et al. (2011) zomes also exhibited antimutagenic effects at
found that the lipidic extract from Iris germanica 1.5, 0.15 and 0.015 mg/plate concentrations.
was able to increase HeLa cell area and adhesion
and augment the formation of actin stress fibres.
This effect was not observed when Ref52 fibro- Antimicrobial Activity
blasts were tested and was not the result of dis-
ruption of microtubules. Further, the increase in The chloroform and ethyl acetate extracts of I.
cell area was Rac1 dependent, and the iris extract germanica rhizomes exhibited bactericidal
led to slight Rac activation. The increase in HeLa activity, while the petroleum ether extract did
cell area in the presence of iris extract was not exhibit any bactericidal, fungicidal and
accompanied by impairment of cell migration insecticidal activities (Orhan et al. 2003 ). It
and arrest of the cell cycle at G1 although there is was also inactive in the brine shrimp toxicity
involvement of Rac1. The active compounds test, whereas it showed significant phytotox-
were found to iridals, a known group of triterpe- icity against the plant Lemna aequinoctialis.
noid. Purified iripallidal was able to increase cell The hexane fraction of the methanol rhizome
area of both HeLa and SW480 cells. extract showed significant inhibitory activity
With respect to antitumour-promoting poten- against Fusarium solani (70 %) and moderate
tial based on anti-inflammatory mechanisms, activity against Trichophyton longifusus
none of the compounds demonstrated significant (50 %) and Microsporum canis (30 %), while
activity in the concentration range tested. Iris the ethyl acetate and chloroform fractions
germanica ceramides irisamides A and B were exhibited moderate activity against the tested
active against mouse lymphoma L5178Y and fungi (Asghar et al. 2009 ). It was observed
cervical cancer HeLa cell lines, but inactive that the growth of bacteria (Pasteurella multo-
against PC12 rat pheochromocytoma (Mohamed cida, Escherichia coli , Bacillus subtilis,
et al. 2013). Irisamides A and B gave 76 % and Staphylococcus aureus) and fungi ( Ganoderma
63 % growth suppression against the L5178Y lucidum, Aspergillus flavus, A. niger,
and HeLa cell lines at a concentration of Alternaria alternata) were inhibited in I. ger-
12.67 mM as well as 68 % and 49 % at a concen- manic a petroleum ether extract (oil) as com-
tration of 3.81 mM, with ED50(s) of 0.91 and pared to the reference standards (rifampicin/
2.40 mM, respectively. The isoflavone iridin S terbinafine) (Asghar et al. 2011). The metha-
was inactive against the three cell lines. Among nolic rhizome extract showed antimicrobial
the compounds isolated from the rhizome, the activity against Staphylococcus aureus,
isoflavone iriskashmirianin A possessed the best Serratia marcescens, Escherichia coli,
cytotoxic activity in Ehrlich’s ascites carcinoma Candida albicans and Aspergillus flavus with
36 Iridaceae
the highest antimicrobial effect against S. IL-2, IFN-gamma and Th2: IL-4 and IL-5 in a
aureus (Ibrahim et al. 2012 ). dose-dependent manner, as studied by the flow
cytometric method. Oral administration of the
isoflavones in Balb/c mice at doses of 0.1–0.8 mg/
Antihyperlipidemic Activity kg per oral dose showed irisolidone to possess
stimulatory activity on T lymphocytes (CD4+
Administration of the ethanolic extract of Iris and CD8+ cells) and Th1: IL-2, IFN-gamma
germanica rhizomes for 10 weeks significantly cytokine production, while irilone acted as an
lowered the lipid components especially the cho- immunosuppressant for T lymphocytes (CD4+
lesterol and triglycerides in rats fed a high-fat and CD8+ cells) and T cell cytokines, namely,
diet (Choudhary et al. 2005) Th1: IL-2, IFN-gamma and Th2: IL-4 and IL-5.
The methylated products of both isoflavones
showed a similar trend to that of their parent
Drug Metabolising Activity compounds, but their activity was drastically
decreased revealing the importance of free phe-
Six isoflavones irisolidone, irisolidone 7-O-α-D- nolic groups for their immunomodulating
glucoside, irigenin, irilone, iriflogenin and iris- activities.
kashmirianin isolated from Iris germanica
rhizomes were shown to be potent inhibitors of
cytochrome P450 1A activity with IC50 values in Antimalarial Activity
the range of 0.25–4.9 μM (Wollenweber et al.
2003). Iridal, a triterpenoidic compound isolated from I.
germanica, exhibited antiplasmodial activity
(Benoit-Vical et al. 2003). The IC50 obtained in-
Anti-inflammatory Activity vitro on human malaria Plasmodium falciparum
chloroquine-resistant and chloroquine-sensitive
Isoflavonoids isolated from I. germanica rhizome strains ranged from 1.8 to 26.0 μg/ml and the
exhibited ant-inflammatory activity determined ED50 in-vivo for P. vincke was about 85 mg/kg/
by a spectrophotometric assay using the activated day by intraperitoneal route. Iridal presented an
human neutrophils (Attar-ur-Rahman et al. antiplasmodial activity similar to that obtained
2003b). The methanolic rhizome extract and the with extracts from the plant Azadirachta indica
isolated flavonoids exhibited potent anti- classically taken as reference in malaria
inflammatory effects by suppressing hind paw phytomedicine
oedema (skin oedema) induced by 4 % formalin
in albino rats (Ibrahim et al. 2012). Irigenin S
showed activity similar to that of Molluscicidal Activity
dexamethasone.
Iris germanica rhizome chloroform extract
showed the highest molluscicidal activity
Immunomodulatory Activity (LC90 = 1.26 mg/l) against Biomphalaria alexan-
drina snails among the tested extracts of the rhi-
Two isoflavones, 5,7-dihydroxy-6,4′-dimetho- zomes (Singab et al. 2006). Fraction B prepared
xyisoflavone (irisolidone) and 5,4′-dihydroxy- from the chloroform extract was the most potent
6,7-methylenedioxyisoflavone (irilone) isolated molluscicide (LC90 = 0.96 mg/l); in addition, it
from Iris germanica, exhibited immunomodula- showed a significant heart rate reduction in the
tory activities (Nazir et al. 2009). They influenced snail after a 6- to 24-h exposure period. It also
the production of T lymphocytes (CD4+ and displayed a significant level of cercaricidal poten-
CD8+ cells) and T cell cytokines, namely, Th1: tial in a time-concentration-dependent manner.
Iris x germanica 37
Traditional Medicinal Uses Abe F, Chem RF, Yamauchi T (1991) Iridals from
Belamcanda chinensis and Iris japonica.
Phytochemistry 30:3379–3382
The root is diuretic, emetic, expectorant and Akashi T, Ishizaki M, Aoki T, Ayabe SI (2005) Isoflavonoid
mildly purgative (Grieve 1971; Usher 1974; production by adventitious-root cultures of Iris ger-
Launert 1981; Chiej 1984). Juice from the root is manica (Iridaceae). Plant Biotechnol 22(3):207–215
38 Iridaceae
Ali AA, El-Emary NA, El-Moghazi MA, Darwish FM, Brenna E, Claudio Fuganti C, Stefano Serra S (2003)
Frahm AW (1983) Three isoflavonoids from Iris ger- From commercial racemic fragrances to odour active
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Nelumbo nucifera
Nelumbo nucifera Gaertner Baladi Bean, Bean Of India, Chinese Water Lily,
East Indian Lotus, Egyptian Bean, Hindu Lotus,
Indian Lotus, Lotus, Lotus Bean, Oriental Lotus,
Synonyms Sacred Lotus, Sacred Water Lotus, Water Lotus
Indonesia: Patma, Tarate (Malay), Tarate, Trate Cattalam, Catti, Cattika, Cattikakkoti,
(Javanese), Tarate (Sundanese) Caturaccakan, Cekkamalar, Cenkalam,
India: Podum (Assamese), Kamal, Kombol, Centamarai,Ceppilai, Ceyyatamarai, Cirukam,
Komal, Padama, Padma, Pudmapudu Cirukam, Citalam, Citampocam, Citampucam,
(Bengali), Ambuj, Kamal, Kamal-Kakri, Cittetan, Civacattipakirtan, Comakkiyam,
Kanwal, Kanval, Lalkamal, Padam (Hindu), Copanam, Cuceyam, Cultalai, Cultalai,
Kamala, Tavare, Tavaribija, Tavarigadde, Curiyanatpu,, Elimanai, Ellimanai, Intai,
Thaavare (Kannada), Bem-Tamara, Iracacupakkoti, Iracacuyam, Iracalam,
Bemtamara, Centamara, Tamara, Ventamara Iraciyam, Iramapiriyam, Iramappiriyam,
(Malayalam), Thambal (Manipuri), Kamal, Iramappiriyam, Iramappiriyam, Irampu,
Kamala (Marathi), Kamal (Oriya), Abja, Iratacanniyakam, Iratikantal, Iratikantan,
Ambhoja, Ambhoruha, Amboruha, Ambuja, Irattacantiyakam, Irattacarorukam,
Ambujanma, Ambupadma, Amburoha, Irattakamalam, Irattakkumutam,
Amlana, Aranala, Aravinda, Arvinda, Irattakokanam, Irattakumutam, Irattalamutam,
Bisakusuma, Bisaprasuna, Drishopadma, Irattamantalam, Irattorpalam, Iravikantam,
Harivetra, Indiralaya, Jalajanma, Kalhara, Irciyam, Jalacam, Kacam, Kamalam,
Kamala, Kamalam, Kamalodbhavamrajah, Kamalini, Kancam, Kancamalar, Kancankoti,
Kanval, Kawar, Kunja, Kusesaya, Kusheshaya, Kantokakkoti, Kantokam, Kantotam, Karkam,
Kutapa, Mahapadma, Mahotpala, Nala, Katirppari, Kauravakkoti, Kauravam,
Nalika, Nalina, Padma, Padmadrajah, Kokanakam, Kokanakam, Kokayam,
Padmam, Pankaj, Pankaja, Pankajam, Makorpalakkoti, Makorpalam, Malunti,
Pankeruha, Pathoja, Pundarika, Pundra, Maraikkoti, Matanalam, Mirinalam,
Pushkara, Puskara, Putaka, Rajiva, Mirunalakkoti, Mirunalam, Mirunali,
Sahasrapatra, Sahsrapatra, Sarasa, Sarasiruha, Mirunarakkoti, Mirunaram, Mirunarram,
Sarojanma, Saroruha, Sarsija, Sarsiruha, Mirutalakkoti, Mirutalam, Muntakakkoti,
Satapatra, Sharada, Sharapadma, Shatapatra, Muntakam, Nalaki, Nalaki, Nalakikkoti,
Shri, Shriparna, Shrivasa, Shuklapadma, Nalalmatu, Nalam, Nalikam, Nalini, Nalitam,
Sitambuja, Sujala, Svetakamala Tamarasa, Nallatamarai, Naticam, Naticam,
Tumarasa, Vanashobhana, Varisoha, Tra Nattuttamarai, Nicakaci,
(Sanskrit), Acaiyapattipam, Acaiyappattiram, NicakacikkotiNirmelcevvanti, Nirmelpaccilai,
Aciyapattiarm, Akiyapattiram, Allakam, Nirnatittamarai, Nirniti, Nirorukakkoti,
Ambal, Ampala, Ampo, Ampocam, Nirorukam, Palututainayakam, Panikam,
Ampocayoni, Ampucam, Ampucani, Panikkanci, Panikkancikkoti, Pankacam,
Ampucanmam, Ampucatam, Ampucini, Pankacatam, Pankacatamakkoti,
Ampunecam, Ampuracam, Ampurecam, Pankacatamam, Pankarukam, Pankatam,
Ampuru, Amputam, Ampuyakkoti, Ampuyam, Pankecam, Pankeram, Panmam, Pannaci,
Ampuyatam, Anikini, Anikini, Appucam, Parparakam, Patmam, Patocam, Patotam,
Arantam, Arappatumam, Arpakantam, Patumakam, Patumanalakkoti, Ponmanai,
Arpatumam, Arukanvakanam, Arunakamalam, Potu, Poykainari, Poykainarinir, Puntaram,
Arunkalaccevvital, Aunikam, Ayamalar, Puntarikam, Putpan, Rattorpalam, Tamaracam,
Aymalar, Caccatam, Calacanam, Calacanmam, Tamaram, Tantamarai, Tantar, Tantulam,
Calacappu, Calacarakkuli, Calakankakkoti, Tanturam, Tarutam, Tevanam, Tikkayam,
Calakankam, Calakarankam, Calarukakkoti, Tirumalarkkoti, Tirumalkompar, Tirumalunti,
Calarukam, Calcam, Calilikam, Cantiram, Totakam, Toyacam, Tunaparicam,
Caracakkoti, Caracam, Caracijam, Sivapputamarai, Tamarai, Thamarai,
Caracirukam, Caravanam, Carocini, Thaamarai, Tamaray, Ventamarai
Carokkam, Carokulam, Catalam, Catapattiram, Vicappicuranam, Vicappiracunam,
Catapatumam, Catippakitakkoti, Catippakitam, Vicaputpam, Vintu, Vintukam Visaputpam,
Nelumbo nucifera 57
Urocanai, Urocani, Urokani, Vacanakkoti, New Guinea, Pakistan, Philippines, Russia (Far
Vacanam, Vanacakkoti, Vanacopanam, East), Sri Lanka, Thailand, Vietnam, Laos,
Vanicam, Vannivannam, Vantuni, Vantunikkoti, Kampuchea and northern Australia.
Varicam, Varicam, Varikam, Vaucikakkoti,
Vaucikam (Tamil), Damara, Erratamara,
Errathaamara, Kalung, Kamalam, Kamalamu, Agroecology
Padmamu, Pundreekamu, Soungadhikamu,
Tamara, Thaamara, Thaamara Puvvu, Thella Lotus is strictly aquatic. It grows wild and is culti-
Thaamara, Thellane Padmanu (Telugu), vated in the mud of shallow, sheltered waters of
Nilufer (Urdu) lakes, ponds and inundated rice fields from sea level
Italian: Fior Di Loto, Giglio De Nilo, Loto to 1500 m elevation in the tropics and subtropics.
d’Egitto, Ninfea d’Egetto
Japanese: Hasu, Hasu-N-Né, Hasu-No-Mi,
Renkon Edible Plant Parts and Uses
Khmer: Chhuk
Korean: Yeongeun, Yon Puri (Rhizome) Lotus is well known as a food and medicinal
Laotian: Bwà plant, and its every part is utilised. All parts of the
Malaysia: Bunga Padam, Bunga Telepok, lotus plant, including flowers, seeds, leaves,
Padema, Seroja, Teratai stems and roots are consumed (Sridhar and Bhat
Nepal: Kamal, Raato Kamal, Raato Thuulo 2007) (Plate 7). In Korea, lotus leaf, root and
Kamal seed are usually consumed as a tea, or in braised
Persian: Beykhneelufer, Nilufer, Nilufu dishes or soups (Ha et al. 2010). Tender rhizomes,
Philippines: Liñgaling (Ibanag), Sukau (Iloko), stems and leaves of lotus are edible and can be
Saua (Maguindanao), Baino (Tagalog) cooked along with other vegetables, soaked in
Portuguese: Flor-De- Lótus, Lótus Do Egipto, syrup or pickled in vinegar (Phillips and Rix
Lótus-Do-Egito, Lótus Índico, Lótus-Sagrado, 1993). Rhizomes have mild flavour and are exten-
Sementes De Lótus sively used in Chinese recipe, while stem is used
Spanish: Habas De Egipto, Loto Sagrado, Rosa in cooking as food and it tastes like beet (Hedrick
Del Nilo, Semillas De Loto, Nelumbio Raiz 1972; Tanaka and Nguyen 2007). Ogle et al.
De Lotus (Root) (2001) reported the use of lotus stem (consisting
Sri Lanka: Nelum (Sinhalese) of 6, 2.4 and 0.2 mg/100 g calcium, iron and zinc,
Taiwan: Lian respectively) as a vegetable used in salads at
Thailand: Bua-Luang, Sattabut, Ubon Vietnam. The seeds can be popped like popcorn,
Tibetan: Pa Dma Dkar Po, Pa Dma Dmar Po, ground into powder and eaten dry or used in
U-Tpa-La bread making. The roasted seeds are good coffee
Vietnam: Hoa Sen, Sen, Lien, Ngau (Tay), Bo substitute and possess saponins. The products of
Bua (Thai), Lin Ngo (Dao) lotus rhizome, such as fresh, salted and boiled
lotus rhizome and lotus rhizome starch, drinks,
teas and lotus seeds, are very popular in the daily
diet (Hu and Skibsted 2007; Zhong et al. 2007).
Origin/Distribution Starches extracted from lotus rhizomes are com-
mercially available in China and consumed in
The species is distributed from the Caspian Sea fast food and as breakfast, traditional confection-
and Iran through Asia to northern Australian – ery and food additives (Zhong et al. 2007). The
China (predominantly in Zhejiang, Jiangsu, Korean traditional lotus liquor (Yunyupju) is
Anhui, Hunan, Hubei), Bhutan, India, Indonesia made from lotus blossom and leaves (Lee et al.
(Java), Japan, Korea, Malaysia, Myanmar, Nepal, 2005). Lotus leaves are used for tea in China
58 Nelumbonaceae
(Zhou et al. 2013a). Lotus leaves are used for tea seed soup dessert’ with longan aril seeds, which
or food like yeon yip bap in Korea (Ahn et al. can be served warm or cold with ice. The Chinese
2013). in China and elsewhere also use lotus seeds as a
The main types of lotus rhizome processed common ingredient in cooling ‘bupin’ mixtures.
products are the salt bloated lotus, deep frozen In India, the seeds after removal of the outer testa
lotus and vacuum-packed lotus (Guo 2009). All is consumed raw, or dried and puffed like
these products are mainly exported to Japan, fol- popcorn.
lowed by South Korean and other southeast Asian The roots (rhizomes) are a food used exten-
countries. Lotus is one of 26 vegetables with sively in China, Korea and Japan and in southeast
largest sale in China (Jiang and Cao 2005). Asia are sold whole or in cut pieces, fresh, frozen
Currently, the lotus herb is becoming more popu- or canned. It has a crunchy texture with sweet-
lar in China as a ‘tea drink’ or as a main ingredi- tangy flavours. The fresh rhizomes are commonly
ent of some herbal formulations (Ye et al. 2014a). sliced and eaten raw or cooked with meat in
Lotus leaf is used as food or beverage in Taiwan soups or fried with salad, prawns, meat, sesame
(Lee et al. 2010). In Sulawesi, the young shoots oil and/or coriander leaves in many Asian cui-
and unexpanded leaves are eaten boiled, and the sines. In Thailand, the young rhizomes are peeled
leaves may be eaten raw (Burkill 1966). The rhi- and eaten fresh in salads, while in China, Japan,
zomes are cooked as food or steam and eaten in Malaysia and elsewhere, the mature rhizomes are
salad. They are also pickled in salt or vinegar. stir-fried, stuffed and deep fried or cooked in
The unripe seeds are eaten raw, boiled or soups. Lotus rhizome is also common in Japanese
roasted, while the ripe seeds are boiled or roasted. ‘bento’ food boxes and ‘nimono’. In Korea, the
Lotus seeds are used in soups, food dishes, sweet- root is used as a vegetable in soups and deep-
meat, cakes, pastries, desserts and nourishing fried, stir-fried and braised dishes. Lotus roots
herbal ‘bupin’ tea. Dried seeds, sometimes called are often pickled with rice vinegar, sugar, chilli
‘lotus nuts,’ must be boiled until softens. The and or garlic. Dried rhizome pieces are also fried
seeds can be crystallised with sugar as part of as chips. A high-quality edible starch is obtained
Chinese New Year sweet offerings, cooked into a from the rhizomes and is used as baby food and
sweet soup called ‘tong sui’ together with dried as a special diet for healing sickness in China.
longan and rock sugar and mashed and cooked The young leaf petioles, after the rough, outer
into sweetened lotus nut paste (‘Lien-yong’) layer has been scraped off, can be found in local
which is used as a filling for Chinese moon cakes markets and are used in preparing salads and
(in mid-Autumn, it is customary to serve ‘moon soups or cooked as vegetable with other meat
cakes’ which have a filling made of lotus seeds dishes in Thailand, Vietnam and Indonesia
and walnuts), ‘daifuku’ (Japanese glutinous rice (Plate 9). The petioles are also slivered (Plate 10).
cake with sweet lotus past as fillings), rice flour In China and Indo-China, flower stamens are
pudding, pastries, buns, bread and sauces. The used to perfume herbal tea called ‘liánhuā cha’ in
seeds are used to prepare ‘tong sui’ (sweet soup). China and trà sen and chè sen in Vietnam.
The seeds can be cooked in soups, usually with Likewise in India, the stamens of the flowers
chicken or beans. An example of the latter is a infused with water are employed to make a fra-
‘red bean and lotus seed soup’ served at banquets grant tea. The dried lotus flowers are used in
for newlyweds, comprising red beans and lotus cooking dishes, such as ‘Mandarin duck and
seeds. Red beans (hong dou) represent strength, lotus flowers’, and the fresh petals are also edi-
while lotus seeds (lian zi) symbolise the newly- ble. In Thailand, young tender leaves are cooked
weds being blessed with a child each year. The and eaten with a savoury sauce, and the flower
soup is also presented at the Chinese New Year’s petals can be dipped in ‘nam prik’ or used as gar-
festival. Other popular and common dishes is the nish. Dried leaves are used for herbal tea in
‘cream lotus seed soup’ which is prepared with China. In Korea, the leaves and petals are used as
crushed pineapple pieces and the ‘sweet lotus a tisane: ‘yeonkkotcha’ made with dried petals of
Nelumbo nucifera 59
white lotus and ‘yeonipcha’ made with leaves. sparsely spinulate (Plates 1, 3). Tepals are cadu-
Lotus leaves are used as wrapper for Chinese cous, pink or white, oblong elliptic to obovate,
‘tamale’, a special Chinese pastry in China. The 5–10 × 3–5 cm and spirally arranged. Stamens are
leaves are used as flavouring and wrapping for numerous, slightly longer than receptacle; fila-
rice preparations in making dim sum and other ment is slender; anther is linear, 1–2 mm; and
sticky, glutinous rice preparations. Lotus leaves connective appendage is clavate, 7 mm long and
are also used for wrapping whole chicken, mari- incurved. Ovary is apocarpous, embedded in an
nated and douched with rice wine, soy sauce, obconical, spongy, accrescent, turbinate 5–10 cm
mushroom and sometimes with some other in diameter receptacle. Its fruit is a nut oblong to
Chinese herbs before the whole entity is encased ovoid, 1.0–2.0 × 7–1.5 cm and glabrous; pericarp
in mud or clay and cooked to give the much- is thick, hardened (bony) and brown (Plates 4, 5).
relished restaurant specialty called ‘beggars Seed is having a reddish brown testa, two yellow-
chicken’ or ‘drunken beggars chicken’. ish-white mealy cotyledons and a stalked green
embryo (Plates 5, 7, 8).
Botany
Nutritive/Medicinal Properties
Nelumbo nucifera is a perennial, rhizomatous,
aquatic herb with the horizontal creeping rhi- Seed/Pod/Receptacle Nutrient/
zomes and roots buried in the mud (Plate 1). The Phytochemicals
rhizome has nodes externally and forms roots at
these nodes, and internally it has aerenchyma, Analyses carried out in the United States reported
which is a parenchyma tissue with large intercel- that raw mature lotus seeds had the following
lular air spaces (Plate 6). The leaves are raised proximate composition (per 100 g value): water,
above the water or floating, on terete, fistulous, 77 g; energy, 89 kcal (372 kJ); protein, 4.13 g;
glabrous or prickly, hollow, 1–2 m long petiole total lipid, 0.53 g; ash, 1.07 g; carbohydrates,
(Plates 9, 10). Leaf’s blade is large, abaxially 17.28 g; Ca, 44 mg; Fe, 0.95 mg; Mg, 56 mg; P,
blue-green, orbicular, peltate, 25–90 cm in diam- 168 mg; K, 367 mg; Na, 1 mg; Zn, 0.28 mg; Cu,
eter, papery, glabrous, glaucous and water repel- 0.094 mg; Mn, 0.621 mg; vitamin C, 0 mg; thia-
lent, having margin entirely (Plates 2, 5). Flowers mine, 0.171 mg; riboflavin, 0.040 mg; niacin,
are large, showy, solitary 10–23 cm in diameter 10.429 mg; pantothenic acid, 0.228 mg; vitamin
on scape longer than petioles, glabrous or B-6, 0.168 mg; total folate, 28 μg; vitamin A,
The ethanol extract of lotus seeds contained tion of diglycerides, monoglycerides, free
more total fatty acids and total phytosterols than fatty acids and free phytosterol (polar compo-
did the ethanol extract of lotus rhizomes. nents) in lotus seeds (g/kg hexane extract) was
Most of the fatty acids were in the triglyceride determined as diglycerides (1,3-dipalmitoylglyc-
ester form in lotus seeds (Zhou et al. 2013b). erol 0.27 g; 1-palmitoyl-2-linoleoylglycerol
Linoleic acid was the most abundant fatty acid 13.17 g; 1-palmitoyl-3-linoleoylglycerol 32.56 g;
compound in lotus seeds and rhizomes. Fatty 1-oleoyl-2-linolenoylglycerol 4.91 g; 1-oleoyl-3-
acids in lotus seeds existed in the glyceride form linolenoylglycerol 14.44 g), monoglycerides
(59.8 %) and to a lesser extent in their free acid (1-palmitoylglycerol 0.80 g; 2-palmitoylglycerol
form (12.0 %). The concentration of fatty acid 12.64 g; 1-linoleoylglycerol 3.60 g,
and phytosterol in the triglyceride and stearyl- 1-oleoylglycerol 2.06 g; 2-behenoylglycerol
fatty acid ester (non-polar components) in lotus 0.26 g), free fatty acids (myristic (C14:0) 0.31 g,
seeds (g/kg hexane extract) was determined as palmitic (C16:0) 43.26 g, linoleic (C18:2)
myristic (C14:0), 0.86 g; palmitic (C16:0), 26.06 g, linolenic (C18:3) 1.26 g, oleic (cis-
59.46 g; linoleic, (C18:2), 176.4 g; linolenic C18:1) 13.20 g, elaidic (trans-C18:1) 0.74 g,
(C18:3), 6.66 g; oleic (cis-C18:1), 43.19 g; stearic (C18:0) 1.10 g, gondoic (C20:1) 0.45 g,
elaidic (trans-C18:1), 3.87 g; stearic (C18:0), arachidic (C20:0) 0.37 g, behenic (C22:0) 2.24 g,
5.40 g; gondoic (C20:1), 1.47 g; arachidic tricosylic (C23:0) 0.38 g, lignoceric acid (C24:0)
(C20:0), 7.77 g; heneicosylic (C21:0), 1.04 g; 0.85 g, total fatty acids 83.91 g) and free phytos-
behenic (C22:0), 28.94 g; tricosylic (C23:0), terols (campesterol 5.25 g, stigmasterol 0.31 g,
1.64 g; lignoceric acid (C24:0), 8.34 g; total fatty β-sitosterol 56.77 g, β-amyrin 1.61 g, total phy-
acids, 345.97 g; campesterol, 1.55 g; isofucosterol, tosterol 63.94 g). Lotus seed oil was found to be
0.12 g; β-sitosterol, 10.57 g; β-amyrin, 5.34 g; an excellent source of crude oil, 3.52 g/100 g dw
and total phytosterol, 17.58 g. The concentration (Anitha and Arunkumar 2012). The oils had an
of diglycerides, monoglycerides, free fatty acids acid value of 26.66 mg KOH, saponification
and free phytosterol (polar components) in lotus value of 210.43 mg KOH, peroxide value of
seeds (g/kg hexane extract) was determined as 10.60 g/100 g, iodine value of 6.00 and free fatty
diglycerides (1,3-dipalmitoylglycerol, 0.27 g; acids of 3.50 g. Crude fat content of lotus seed oil
1-palmitoyl-2-linoleoylglycerol, 13.17 g; hexane extract was 38.6 %, and fatty acid methyl
1-palmitoyl-3-linoleoylglycerol, 32.56 g; 1-oleoyl- esters found included capric acid methyl ester
2-linolenoylglycerol, 4.91 g; 1-oleoyl-3- (C10:0), lauric acid methyl ester (C12:0), tri-
linolenoylglycerol, 14.44 g), monoglycerides decanoic acid methyl ester (C13:0), palmitic acid
(1-palmitoylglycerol, 0.80 g; 2-palmitoylglycerol, methyl ester (C16:0), pentadecanoic acid methyl
12.64 g; 1-linoleoylglycerol, 3.60 g; ester (C15:0), cis-10-pentadecanoic acid methyl
1-oleoylglycerol, 2.06 g; 2-behenoylglycerol, ester (C15:1), palmitoleic acid methyl ester
0.26 g), free fatty acids (myristic (C14:0), 0.31 g; (16:1), cis-10-heptadecanoic acid methyl ester
palmitic (C16:0), 43.26 g; linoleic (C18:2), (C17:1) and nonadecanoic acid methyl ester
26.06 g; linolenic (C18:3), 1.26 g; oleic (cis- (C19:1).
C18:1), 13.20 g; elaidic (trans-C18:1), 0.74 g; Seed starch grain was oval and contained
stearic (C18:0), 1.10 g; gondoic (C20:1), 0.45 g; higher amylase content starch and gelatinisation
arachidic (C20:0), 0.37 g; behenic (C22:0), temperature than rhizome (Man et al. 2012). Seed
2.24 g; tricosylic (C23:0), 0.38 g; lignoceric acid starches exhibited an A-type X-ray diffraction
(C24:0), 0.85 g; total fatty acids, 83.91 g) and pattern with higher crystalline degree. Seed
free phytosterols (campesterol, 5.25 g; stigmas- starch had lower acid hydrolysis and higher por-
terol, 0.31 g; β-sitosterol, 56.77 g; β-amyrin, cine pancreatic α-amylase hydrolysis than rhi-
1.61 g; total phytosterol, 63.94 g). The concentra- zome starch. A novel lotus plumule polysaccharide
64 Nelumbonaceae
The procyanidins of lotus seedpod were heat-induced annexin, NnANN1, was identified
extracted with aqueous acetone with a purity of from lotus embryonic axes that was found to play
>98 % (Lin et al. 2005). The main molecular an important role in seed thermotolerance and
weight distribution of procyanidins ranged from germination vigour. Compared to the wild-type
291 to 1155, with M + H peak values of 291.1, seeds, transgenic seeds ectopically expressing
579.2, 731.2, 867.2, 1019.4 and 1155.3, respec- NnANN1 exhibited improved resistance to accel-
tively, indicating the extract contained mono- erated ageing treatment. Also transgenic seeds
mers, dimers and tetramers of procyanidins, in showed enhanced peroxidase activities, accom-
which the amounts of dimers were greatest, and panied with reduced lipid peroxidation and
catechin and epicatechin were the base units. The reduced ROS release levels compared to the
procyanidins of lotus seedpod were extracted wild-type seeds. Studies by Shen-Miller et al.
with aqueous acetone with a purity of >98 % (Lin (2013) showed that in proteins of N. nucifera
et al. 2005). The main molecular weight distribu- fruit that were able to withstand heating, 31 % of
tion of procyanidins ranged from 291 to 1155, which remained soluble in the 110 °C-treated
with M + H peak values of 291.1, 579.2, 731.2, embryo-axis of a 549-yr-old fruit and 76 %
867.2, 1019.4 and 1155.3, respectively, indicat- retained fluidity in its cotyledons. At 110 °C for
ing the extract contained monomers, dimers and 10 min, 22.6 μg/mg of thermal proteins were
tetramers of procyanidins, in which the amounts recovered from the embryo-axis of a 549-year-
of dimers were greatest, and catechin and epicat- old lotus seed versus 7.1 μg/mg from modern
echin were the base units. Five flavonol glyco- seeds; 11.1 μg/mg were recovered from the coty-
sides, including hyperoside, isoquercitrin, ledons of a 549-year-old seed versus 21.9 μg/mg
quercetin-3-O-ß-d-glucuronide, isorhamnetin-3- from the cotyledons of modern seeds. The amino-
O-ß-d-galactoside and syringetin-3-O-ß-d-- acid sequences of 11 “thermal proteins” (soluble
glucoside, were determined in lotus receptacle at 100 °C) of modern Nelumbo embryo-axes and
with recoveries in the range of 98.31–100.32 % cotyledons were identified as dehydrin (gluta-
(Wu et al. 2013). The oligomeric and polymeric mine, lysine), 55 % total amino acid; HSP80
fraction of the 60 % aqueous methanol lotus seed (lysine/glutamine, asparagine), 38 %; CPN60
pod extract had a mean degree of polymerisation (lysine, glutamine), 33 %; vicilin (glutamine,
of 3.2 and 15.4, respectively, and consisted of lysine, arginine/asparagine), 30 %; EF-1α (lysine,
(+)-catechin (m/z 289), gallocatechin or epigal- asparagine, glutamine), 29 %; met-synthase (glu-
locatechin (m/z 305), quercetin glycoside (m/z tamine, lysine, asparagine), 36 %; 1-CysPRX
463), quercetin glucuronide (m/z 477), procyani- (lysine, asparagine), 26 %; ENO1 (lysine, gluta-
din dimers (m/z 577.1), proanthocyanidin dimer mine), 24 %; CPN20 (lysine, glutamine, aspara-
gallate (m/z 593.3), prodelphinidin dimers (m/z gine), 22 %; PIMT (glutamine, lysine,
609.1), procyanidin trimers (m/z 865.1), etc asparagine), 21 %; and CuZn-SOD (asparagine,
(Xiao et al. 2012). Quercetin glucuronide was lysine, glutamine), 18 %. The 11 Nelumbo
further purified and identified as quercetin thermal proteins studied could be grouped into
-3-O-β-glucuronide. A flavonoid quercetin [2- four functionally defined categories: antioxidant
(3,4-dihydroxy)-3, 5, 7-trihydroxy-4H-1-benzo- and membrane maintenance (CuZn-SOD,1-
pyran-4-one] was isolated from lotus dried recep- CysPRX, dehydrin); chaperonins, chaperone and
tacle (Ishida et al. 1988). Five flavonol glycosides, stress proteins (Cpn20, Cpn60, HSP80, EF-1α);
namely, hyperoside, isoquercitrin, quercetin- defence, food and repair (vicilin, Met-Synthase,
3-O-β-d-glucuronide, isorhamnetin-3-O-β-d- PIMT); and anaerobic glycolytic protein
galactoside and syringetin-3-O-β-d-glucoside, (Enolase1 (ENO1)). In addition, 30 other heat-
were isolated from lotus receptacle (Wu et al. stable proteins of N. nucifera cv. China Antique
2012). were identified: actin 12, aldose reductase, aller-
Lotus seeds were found to contain annexins, gen Ara h1 clone, nep1 aspartic proteinase
multifunctional proteins (Chu et al. 2012). A nepenthesin 1, chaperone protein yajL, embry-
Nelumbo nucifera 67
onic protein DC-8, pistil-specific extensin-like drates, 17.23 g; total dietary fibre, 4.9 g; Ca,
protein, FBA fructose biphosphate aldolase 45 mg; Fe, 1.16 g; Mg, 23 mg; P, 100 mg; K,
(cytosol enzyme), 11S globulin β, FA02 13S 556 mg; Na, 40 mg; Zn, 0.39 mg; Cu, 0.257 mg;
globulin seed storage protein 1, GluA1 glutelin Mn, 0.261 mg; Se, 0.7 μg; vitamin C, 44 mg;
type A1, GluB1 glutelin type B1, GAPC thiamine, 0.160 mg; riboflavin, 0.220 mg; niacin,
glyceraldehye-3-phosphate dehydrogenase 0.400 mg; pantothenic acid, 0.377 mg; vitamin
(cytosol), HSP18.2 18.2 kDa Class 1 heat shock B-6, 0.258 mg; total folate, 13 μg; total saturated
protein and HSP70; late embryonic abundant fatty acids, 0.030 g; 16:0 (palmitic acid), 0.028 g;
(LEA) proteins Dc3, Dc 8 and Dc 34; and 18:0 (stearic acid), 0.001 g; total monounsatu-
Legumin A (LEGA), Legumin B (LEGB), low rated fatty acids, 0.020 g; 16:1 undifferentiated
temperature-induced 65 kDA (LT65) protein, (palmitoleic acid), 0.002 g; 18:1 undifferentiated
malate dehydrogenase (MDH), 5-methyltetrahyd (oleic acid), 0.014 g; 20:1 (eicosenoic acid),
ropteroyl-triglutamate-homocysteine methyl- 0.002 g; total polyunsaturated fatty acids,
transferase (metE), early nodulin-like protein 0.020 g; 18:2 undifferentiated (linoleic acid),
1(Atg25060), protein notum homolog (NOTUM), 0.014 g; 18:3 undifferentiated (linolenic acid),
pectiesterase 2 (PECS-2.1), PCKR 1 peptidyl- 0.02706 g; tryptophan, 0.020 g; threonine,
prolyl cis-trans isomerise, peroxidase 12 0.051 g; isoleucine, 0.054 g; leucine, 0.069 g;
(PER12), phosphoglycerate kinase (cytosol) and lysine, 0.094 g; methionine, 0.022 g; cystine,
tricohyalin and triose-phosphate isomerase 0.022 g; phenylalanine, 0.047 g; tyrosine,
(TCHH). 0.029 g; valine, 0.055 g; arginine, 0.088 g; histi-
The main constituents in the essential oils dine, 0.038 g; alanine, 0.054 g; aspartic acid,
from the various rhizome and seed samples were 0.369 g; glutamic acid, 0.139 g; glycine, 0.156 g;
l-(+)-ascorbic acid 2,6-dihexadecanoate proline, 0.136 g; and serine, 0.060 g (USDA-
(0–33.5 %), trans-phytol (5.1–24.1 %), hexahy- ARS 2014).
drofarnesyl acetone (5.6–15.3 %), pentadecyl The nutrient composition of the rhizome per
acrylate (2.2–12.4 %), geranyl acetone (1.9– 100 g edible portion (CINE 2007) was reported
8.0 %) and β-ionone (0–8.0 %) (Huang et al. as follows: moisture, 80.5 g; energy, 75 kcal; pro-
2010a). The rhizome lotus and seed lotus sam- tein, 1.9 g; fat, 0.2 g; carbohydrate, 16.4 g; fibre,
ples were clustered into separate groups by hier- 1.2 g; ash, 1 g; vitamin A, 3 μg RE (1.5 μg RAE);
archical cluster analysis according to the total carotene, 20 μg; vitamin C, 44 mg; Zn,
composition of the corresponding essential oils. 0.23 mg; Fe, 1.4 mg; and Ca, 39 mg. Another
No significant relationship was found between study reported that the rhizomes have thiamine,
essential oil composition and geographical distri- 0.16 mg; riboflavin, 0.22 mg; niacin, 0.4 mg;
bution of the 11 populations. vitamin C, 44 mg; calcium, 45 mg; phosphorus,
Young chloroplasts were found in shoots of 100 mg; iron, 1.6 mg; potassium, 730 mg;
mature dry lotus seeds, and they gave rise to sodium, 40 mg; magnesium, 25 mg; zinc, 0.2 mg;
mature chloroplasts during germination, even in protein, 2.6 g; carbohydrate, 14.7 g; and fat, 0.1 g
darkness (Ushimaru et al. 2003). These shoots (Howard et al, 1962). Korean white lotus culti-
contained chlorophyll and chlorophyll-binding vars (Inchisa, Muan, Garam and Chungyang) and
proteins CP1 and LHCP. rhizomes were reported to possess high amounts
of bioactive compounds: total phenols, between
7.95 and 4.21 mg of gallic acid equivalents
Rhizome Nutrients/Phytochemicals (GAE)/g dry weight (DW); ascorbic acid,
between 15.8 and 22.3 mg of ascorbic acid/g
The nutrient composition of raw rhizome per DW; and amino acids, between 15.05 % and
100 g edible portion was reported as follows: 16.62 0 % DW (Park et al. 2009). Tryptophan was
water, 79.10 g; energy, 74 kcal (311 kJ); protein, isolated from the aqueous extract of lotus rhi-
2.60 g; total lipid, 0.10 g; ash, 0.97 g; carbohy- zome (Jiang et al. 2010).
68 Nelumbonaceae
The ethanol extract of lotus rhizomes con- respectively, while least gelation concentration,
tained less total fatty acids and total phytosterols foaming volume increase, foaming stability,
than did the ethanol extract of lotus seeds (Zhao emulsifying capacity and emulsion stability were
et al. 2013a). The concentration of fatty acid and 18.0 %, 5.23 %, 4.97 %, 48.93 % and 96.43 %,
phytosterol in the triglyceride and stearyl-fatty respectively. The lotus rhizome was found to be a
acid ester (non-polar components) in lotus rhi- poor source of crude oil (2.68 g/100 g dry weight)
zomes (g/kg hexane extract) was determined as with the following physicochemical properties:
follows: palmitic (C16:0), 43.64 g; linoleic crude fat (2.68 g/100 g DW), acid value (16.66 mg
(C18:2), 53.68 g; linolenic (C18:3), 7.91 g: KOH), saponification value (110.43 mg KOH)
oleic (cis-C18:1), 14.27 g: elaidic (trans- and unsaponifiable matter (0.026 g/100 g oil).
C18:1), 1.17 g; stearic (C18:0), 8.34 g; ara- The amylose content was the highest
chidic (C20:0), 0.43 g; behenic (C22:0), 1.56 g; (30.61 %) in lotus starch (Zhiong et al. 2007).
and total fatty acids, 131.01 g. The concentra- The average particle size (diameter) was 50.27,
tion of diglycerides, monoglycerides, free fatty 24.08 and 38.97 μm for lotus, kudzu and corn
acids and free phytosterol (polar components) starches, respectively. Lotus starch exhibited a
in lotus rhizomes (g/kg hexane extract) was B-type X-ray diffraction pattern and kudzu starch
determined as follows: diglycerides (0.27 g exhibited a C-type pattern. Kudzu starch was
1-palmitoyl-2-linoleoylglycerol and 0.45 g characterised by a maximum viscosity immedi-
1-palmitoyl-3-linoleoylglycerol), monoglycer- ately followed by a sharp decrease in viscosity,
ides (8.13 g 2-palmitoylglycerol and 7.11 g while the lotus starch was characterised by a pla-
1-linoleoylglycerol), free fatty acids (myristic teau when the maximum viscosity was reached.
(C14:0), 0.11 g; palmitic (C16:0), 6.80 g; linoleic Lotus rhizome starches had small oval gran-
(C18:2), 5.21 g; linolenic (C18:3), 1.51 g; oleic ules and large elongated granules (Man et al.
(cis-C18:1), 1.51 g; elaidic (trans-C18:1), 0.15 g; 2012). Rhizome starches had higher swelling
stearic (C18:0), 0.41 g; gondoic (C20:1), 0.02 g; power, lower amylase content and gelatinisation
arachidic (C20:0), 0.09 g; heneicosylic (C21:0), temperature than seed starch. Rhizome starches
0.02 g; behenic (C22:0), 0.34 g; tricosylic showed a C-type pattern which changed from C
(C23:0), 0.08 g; lignoceric acid (C24:0), 0.45 g; to A type with gradually increasing crystalline
and total fatty acids, 83.9116.02 g) and free phy- degree during acid hydrolysis. The degree of
tosterols (campesterol, 4.51 g; isofucosterol, order in starch external region was higher in rhi-
0.65 g; β-sitosterol, 76.27 g; β-amyrin, 7.34 g; zome than in seed. The external region structure
α-amyrin, 7.78 g; lanosterol, 12.68 g; and total of rhizome starches became more ordered during
phytosterol, 109.22 g). acid hydrolysis. Rhizome starches had higher
The proximate composition of lotus rhizome rate of acid hydrolysis and lower rate of porcine
flour (g/100 g flour) was determined as ash pancreatic α-amylase hydrolysis than seed
(1.10 g), the total nitrogen (1.36 g), total protein starches. Starch in rhizome of lotus cultivar
(8.48 g), water-soluble proteins (1.23 g), salt- Meirenhong exhibited C-type X-ray diffraction
soluble proteins (5.73 g), total salt-soluble pro- pattern, while starch in rhizome of cultivar
teins (6.064 g), total sugar (19.08 g), reducing Wawalian showed A-type pattern (Yu et al.
sugars (0.168 g), non-reducing sugars (18.87 g) 2013a). Starch granules in cv. Meirenhong
and free amino acids (0.78 g) (Shad et al. 2011). showed oval-shaped granules, while starch gran-
The temperature-dependent behaviour of solubil- ules in cv. Wawalian were elongated and oval in
ity and swelling capacity of the rhizome flour shape with relatively large size. Gelatinisation
showed a linear increase (1.2–13.84 %) in solu- temperatures of starch in Meirenhong and
bility but an exponential increase in swelling Wawalian were 330.5 and 342.4 K, respectively,
capacity with a gradual increase in the tempera- and the gelatinisation temperature range of
ture (40–100 °C). Water absorption and oil hold- Meirenhong was significantly wider than that of
ing capacities were found to be 2.56 and 2.03 %, Wawalian. Starch in rhizome of cv. Meirenhong
Nelumbo nucifera 69
showed lower pasting temperature, lower hot and extracts yielded alkaloids, steroids, reducing sug-
cool viscosities, lower setback and higher peak ars and saponins (Mukherjee et al. 1998). The
viscosity and breakdown than those of cv. detection and quantification limitations of cate-
Wawalian in rapid viscosity analyser pasting pro- chin were 23 ng and 50 ng, respectively, in lotus
files at 6 % starch concentration. Amylose mole- rhizome (Yan et al. 2009). The catechin level was
cules in lotus rhizome starch were large, with 0.0025 % in the lotus rhizome and 0.011 % in the
number-average and weight-average degrees of knot of the lotus rhizome. (±)-Gallocatechin and
polymerisation (DPs) of 4170 and 8040, respec- (–) catechin plus LB2 was a polysaccharide-
tively, number-average chain length (͞͞CLn) of 540 protein complex with a molecular mass of
and apparent DP distribution range of 520– 18.8 kDa isolated from lotus rhizome (Jiang et al.
42,000 (Suzuki et al. 1992). The ͞CLn of amylo- 2011). LB2 was identified as a polysaccharide
pectin was 22.3, and its distribution by sulphate containing α/β-pyranose and α-furanose.
gel-exclusion HPLC showed three peaks, at DP It was composed of mannose, rhamnose, glucose,
14, 42 and 1900, and a shoulder at DP 63. The galactose and xylose with a molar ratio 2:8:7:8:1.
amylose content was 15.9 % determined by The percentage of recovery of betulinic acid
iodine affinity and 17.4 % determined by blue from lotus rhizome by HPTLC was found to be
value. The onset of gelatinisation determined by 98.36 % (Mukherjee et al. 2010c). The limit of
photopastegraphy was at 58.5 °C. The amylo- detection and limit of quantification of betulinic
gram of the starch resembled that of tapioca acid were detected to be 0.4 and 2.30 μg per spot.
starch. The retrogradation tendency of lotus From lotus rhizome, a new ursane triterpenoid
starch was slower than that of potato, lily and ester, urs-12-en-3β-O-9E,12E-octadecadienoate,
kudzu starches but faster than that of tapioca was identified along with the isolation of seven
starch. During gelatinisation of lotus rhizome known compounds (palmitic acid, linoleic acid,
starch, the crystallinity changed from C type to A 9E,12E,15E-octadecatrienoic acid, α-amyrin,
type via CA type and finally became amorphous β-sitosterol, betulinic acid and β-sitosterol-3-O-
structure (Cai et al. 2014). The amylose content, glucoside) (Chaudhuri and Singh 2009) and a
crystal degree, helix content, ratio of new triterpenoid, 2α,24-diacetoxy-3β-
1045/1022 cm−1 and peak intensity of crystalline hydroxyolean-12-en-28-oic acid, along with four
lamellae of gelatinising starch significantly known compounds (hyptatic acid-A (2α,3β,24-
decreased after 70 °C. The amorphous content trihydroxyolean-12-en-28-oic acid), maslinic
and ratio of 1022/995 cm−1 increased after acid (2α,3β-dihydroxyoleane-12-ene-28-oic
70 °C. The study elucidated that B-type allo- acid), betulin and lupeol) (Chaudhuri and Singh
morph was mainly arranged in the distal region 2013).
of eccentric hilum, A-type allomorph was mainly Fifteen compounds were isolated from the
located in the periphery of hilum end, and the total alkaloid extract of lotus stem (rhizome) and
centre of the starch granule was a mixed distribu- identified as asimilobine, isococlaurine,
tion of A- and B-type allomorphs. Lotus seed- N-acetylnorarmepavine, crykonisine, velucryp-
resistant starch (LRS), a type of retrograded tine, pycnarrhine, liriodenine, nuciferine, nor-
starch, is commonly known as resistant starch nuciferine, armepavine, N-methylasimilobine,
type 3 (RS3) (Zhang et al. 2014). Compared with coclaurine, N-norarmepavine (13),
native starch and high amylose maize starch N-methylcoclaurine and lysicamine (Duan et al.
(HAMS), LRS lacked the polarisation cross, and 2013). Astragalin, rutin, isoquercetin, nuciferine,
the irregularly shaped LRS granules had a dauricine, isoliensinine and neferine were identi-
rougher surface, B-type crystal structure and fied in lotus rhizomes (Zhao et al. 2014).
greater level of molecular order. High-quality RNA was recovered from five
The petroleum ether extract of lotus rhizomes different tissues of lotus as assessed by polyphe-
yielded steroids; the chloroform extract yielded nol oxidase (PPO) gene expression profiles
alkaloids and steroids; the methanol and water (Zhang et al. 2013b). The relative expression lev-
70 Nelumbonaceae
els of the PPO gene based on RTPCR in five tis- zome (Tomita et al. 1961c). Roemerine and
sues of lotus were rhizome buds (2.66), young nuciferine were isolated as the tertiary, non-
leaves (2.42), fresh cut rhizome (2.02), petals phenolic bases, and a new base, nornuciferine, as
(1.80) and petiole (1.65), using housekeeping the tertiary phenolic base, from the leaves.
gene β-actin as an internal control. The ethanol Roemerine and nornuciferine were isolated from
extracts of lotus seeds from Vietnam (Ho Chi the petiole but not from nuciferine. The results
Minh City), raw rhizomes from Korea (Siheung) indicated that the main alkaloids of lotus are all
and dried rhizomes from Japan (Nigata) had the tertiary bases of the aporphine type. Roemerine,
greatest specific flavonoid content (Zhao et al. nuciferine and nornuciferine, a new aporphine-
2014). The ethanol extracts of seeds from China type tertiary phenolic base, were isolated from
(Hubei), raw rhizomes from Japan (Nigata) and the leaves and petioles of domestic lotus (Tomita
dried rhizomes from Korea (Siheung) had the et al. 1961a). Besides the isolation of phenolic
greatest specific alkaloid content. bases, nuciferine, roemerine and nornuciferine,
another base, dl-armepavine, was isolated from
the leaves (Tomita et al. 1961b). Aporphine-type
Leaf Nutrients/Phytochemicals alkaloid bases, roemerine, and nornuciferine,
and benzyl-tetrahydro-isoquinoline-type base,
Proximate nutrient composition of lotus leaf (g armepavine, from the leaves, and roemerine and
DW/100 g) was reported as moisture, 1.65 g; nornuciferine from the leaf petiole were isolated
crude protein, 14.47 g; crude lipid, 2.29 g; crude from an ancient lotus cultivar Ohga-hasu (Tomita
ash, 9.36 g; carbohydrate, 10.23 g; chemicals and Frurukawa 1962). Alkaloids contained in the
(mg DW)/100 g K, 1345.25 mg; Ca, 1069.51 mg; so-called primitive lotus ‘gen-shi-hasu’ were
P, 293.27 mg; Mg, 261.48 mg; Na, 12.4 mg; Fe, similar to domestic lotus: roemerine, nuciferine,
4.32 mg; Zn, 1.76 mg; Cu, 0.26 mg; sucrose, nornuciferine and dl-armepavine (Kunitomo
1646.56 mg; fructose, 210.70 mg; glucose, et al. 1964). Cleavage reaction of nuciferine and
110.14 mg; vitamin A, 714.61 mg; vitamin C, nornuciferine by metallic sodium in liquid
679.59 mg; vitamin B1, 4.10 mg; and vitamin ammonia afforded 2-methoxyaporphine and
B2, 3.67 mg (Ha et al. 2010). 2-hyrdoxyaporphine, respectively (Kunimoto
Scanning electron microscopy and chemical and Kamimura 1964). Tertiary alkaloids
analysis of the chloroform extract of lotus leaves anonaine, pronuciferine, N-nornuciferine, liri-
showed that the wax was composed of a mixture odenine and d-methylcoclaurine were isolated
of aliphatic compounds, principally nonacosanol from the leaves besides the known roemerine,
and nonacosanediols (Koch et al. 2006). Analysis nuciferine, O-nornuciferine and dl-armepavine
of gas chromatography spectra of lotus leaf (Kunitomo et al. 1970). The structure of nor-
waxes showed a much lower proportion of the nuciferine, an aporphine-type tertiary phenolic
secondary alcohol nonacosan-10-ol (16.2 % by base isolated from the domestic lotus, was proven to
weight) compared with nonacosanediols be dl-1-methoxy-2-hydroxyaporphine (Kunitomo
(64.7 %). Gas chromatographic analysis of the et al. 1971). Four new alkaloids, dehydroro-
extracted leaf waxes revealed nonacosan-10-ol emerine, dehydronuciferine, dehydroanonaine
(16.2 %), triacontan-7-ol (2.4 %), nonacosane-4, and N-methylisococlaurine, were isolated from
10-diol (18.6 %), nonacosine-5, 10-diol (34.1 %), N. nucifera leaves, besides the known roemerine,
nonacosane-10, 13-diol (12.0 %), hentriacon- nuciferine, anonaine, pronuciferine,
tane- 12, 15-diol (1.8 %), tritriacontane-9, 10-diol N-nornuciferine, nornuciferine, amepavine and
(0.7 %) and octadecanoic acid (0.7 %). N-methylcoclaurine (Kunitomo et al. 1973).
The first alkaloid isolated from lotus leaves Asimilobine and lirinidine, serotoninergic recep-
was nuciferine (Arthur and Cheung 1959). tor antagonists, were isolated from lotus leaves
Alkaloids were found in lotus leaves, petiole and (Shoji et al. 1987). Four alkaloids, i.e., 2-hydroxy-
cotyledons in the fruit but not in the edible rhi- 1-methoxyaporphine, pronuciferine, nuciferine
Nelumbo nucifera 71
and roemerine, were identified in Nelumbo nuciferine and nornuciferine, was isolated from
nucifera and its alkaloid fraction by HPLC (Wang Nelumbo nucifera leaves (Yang et al. 2012). The
et al. 2008). The linear ranges of 2-hydroxy-1- following major alkaloids (−)-nuciferine,
methoxyaporphine, pronuciferine, nuciferine and (−)-nornuciferine, (−)-caaverine, (−)-armepavine,
roemerine were 0.110–0.658 μg (R2 = 0.9995), (+)-norarmepavine, (+)-isoliensinine and
0.0210–0.126 μg (R2 = 0.9995), 0.103–0.618 μg (+)-pronuciferine were found in Nelumbo
(R2 = 0.9998) and 0.085 6–0.514 μg (R2 = 0.9995), nucifera leaves (Do et al. 2013). In all samples,
with the average recoveries of 101.5 %, 99.14 %, (−)-nuciferine (0.34–0.63 %), (−)-armepavine
99.21 % and 98.41 % for the alkaloid fraction of (0.13–0.20 %) and (+)-isoliensinine (0.06–
N. nucifera and 99.53 %, 100.5 %, 97.51 % 0.19 %) were the most dominant alkaloids. The
and 100.1 % for N. nucifera, respectively. total alkaloid content varied from 0.72 to 1.41 %.
(+)-1(R)-Coclaurine, (–)-1(S)-norcoclaurine, From the methanolic extracts of lotus flower buds
liensinine, neferine, isoliensinine and nuciferine and leaves, a new alkaloid, N-methylasimilobine
together with quercetin 3-O-β-d-glucuronide (4) N-oxide, was isolated together with 11 benzyliso-
were isolated from lotus leaves (Kashiwada et al. quinoline alkaloids (Nakamura et al. 2013).
2005). N-Nornuciferine, O-nornuciferine, nucif- 2-Hydroxy-1-methoxyaporphin (2H1M), an
erine and roemerine were identified in lotus alkaloid, was isolated from lotus leaves (Chen
leaves (Luo et al. 2005). From 4.0 g of the crude et al. 2012c). Major anti-inflammatory com-
lotus leaf extract, 120 mg N-nornuciferine, pounds identified from lotus leaves were nucifer-
1020 mg nuciferine and 96 mg roemerine were ine, O-nornuciferine, hirsutrin, hyperin,
obtained each with a purity of over 98 % (Zheng luteoloside and roemerine (Zhou et al. 2013a).
et al. 2010). Three alkaloids N-nornuciferine, From lotus leaves, liriodenine, lysicamine,
O-nornuciferine and nuciferine were isolated (−)-anonaine, (−)-asimilobine, (−)-caaverine,
from lotus leaf (Ma et al. 2010). 500 mg of crude (−)- N -methylasimilobine, (−)-nuciferine,
lotus leaf extract furnished 7.4 mg of (−)-nornuciferine, (−)-roemerine, cepharadione
N-demethylarmepavine, 45.3 mg of nuciferine B and 7-hydroxydehydronuciferine were isolated
and 26.6 mg of roemerine with purities of 90 %, (Lin et al. 2014). 100 mg of crude lotus leaf
92 % and 96 %, respectively (Xu et al. 2011b). extracts yielded 6.3 mg of 2-hydroxy-1-
The compounds isolated from lotus leaves methoxyaporphine (95.1 % purity), 1.1 mg of
included seven aporphines ((–)-nuciferine, pronuciferine (96.8 % purity), 8.5 mg of nucifer-
(–)-nornuciferine, (–)-asimilobine,(–)-N- ine (98.9 % purity) and 2.7 mg of roemerine
methylasimilobine, (–)-caaverine, (–)-anonaine (97.4 %) (Ma et al. 2014).
(6) and (–)-roemerine), two oxoaporphines (lysi- Quercetin, myricetin, kaempferol
camine and liriodenine), one dioxoaporphine 3-O-glucoside, quercetin 3-O-glucoside and
(cepharadione B), one dehydroaporphine luteolin 7-O-glucoside were isolated from aer-
(–7-hydroxydehydronuciferine), two steroids ial parts of N. nucifera (Wassel et al. 1996).
(β-sitostenone and stigmasta-4,22-dien-3-one) From lotus leaf extract, the following com-
and two chlorophyll (pheophytin-a and aristo- pounds were isolated: a new megastigmane,
phyll-C) (Wang et al. 2011). (–)-caaverine, ceph- nelumnucifoside, together with 12 megastig-
aradione B and 7-hydroxydehydronuciferine manes ((E)-3-hydroxymegastigm-7-en-9-one,
were reported for the first time in lotus. (−) –boscialin, (+) – dehydrovomifoliol, vomi-
Liriodenine in lotus leaves was extracted and foliol, 3-oxo-retro-α-ionol I, byzantionoside A,
quantified using ethanol and water-soluble sol- 5,6-epoxy-3- hydroxy-7-megastigmen-9-one,
vents; liriodenine content obtained was 4.8 mg% annuionone D, icariside B2, grasshopper
by both methods (Mongkolrat et al. 2012). ketone, 3S, 5R-dihydroxy-6S, 7-megastigma-
N-methylasimilobine, a newly found potent dien-9-one and (+)-epiloliolide); a new eudes-
acetylcholinesterase (AChE) inhibitor, along mane sesquiterpene, eudesnelumboside; eight
with two weakly active aporphine alkaloids, alkaloids ((6R, 6aR)-roemerine-Nβ-oxide,
72 Nelumbonaceae
oil was 7.18 g/kg; carotenoids, 377.86 mg/kg; galactoside (f13), kaempferol 3-O-robinobioside
squalene, 84.94 mg/kg; and sterols, 490.90 mg/ (f14), isorhamnetin 3-O-rutinoside (f15), kaemp-
kg dry bee pollen. ferol 3-O-β-d-glcuopyranoside (f16), kaempferol
3-O-glucuronide (f17), isorhamnetin 3-O-β-d-
glucopyranoside (f18), isorhamnetin 3-O-glucu-
Aerial Plant Parts Phytochemicals ronide (f19), syringetin 3-O-glucuronide (f20),
apigenin di-C-glucoside (f21), 6-C-glucosyl-8-C-
Alkaloids were found in the leaves, petiole and pentosyl luteolin (f22), 6-C-pentosyl-8-C-glucosyl
cotyledons in the fruit but not in the edible rhi- luteolin (f23), 6-C-glucosyl-8-C-xylosyl api-
zome (Tomita et al. 1961c). All the alkaloids genin (f24), 6-C-xylosyl-8-C-glucosyl api-
found were in a free state. Roemerine and nucif- genin (f25), 6-C-glucosyl-8-C-arabinosyl api-
erine were isolated as the tertiary, non-phenolic genin (schaftoside) (f26), 6-C-arabinosyl-8-C-
bases, and a new base, nornuciferine, as the ter- glucosyl apigenin (isoschaftoside) (f27), quer-
tiary phenolic base, from the leaves. Roemerine cetin 3-O-neohesperidoside (f28), 6-C-gluco-
and nornuciferine were isolated from the petiole syl-8-C-rhamnosyl apigenin (f29), luteolin
but not nuciferine. After methanolic extraction, 7-O-rutinoside (f30), 6-C-rhamnosyl-8-C-glu-
the phenolic content of each part of the plant, cosyl apigenin (f31), isorhamnetin 3-O-neohes-
determined in terms of gallic acid equivalents per peridoside (f32) and diosmetin 7-O-rutinoside
g of dried extract (GAE, mg/g extract), was found (f33). Flower petals and stamens possessed the
to follow the order of leaves (177.7) > de- highest amounts of kaempferol glycoside, spe-
embryonated seeds (92.7) > stamens (83.4) > cially f11 and f17. Additionally f14 and f16 were
embryos (41.0) > rhizomes (21.6). For the flavo- the predominant compounds in petals. The petals
noid contents, determined as mg of (+)-catechin contained f1, f4, f5, f7, f8, f10, f11, f14, f15, f16,
equivalent per g of dried extract (CE, mg/g f17, f18 and f19. The stamens contained f1, f4,
extract), the order was leaves (125.6) > de- f5, f7, f8, f10, f11, f12, f14, f15, f16, f17, f18 and
embryonated seeds (82.9) > stamens (50.3) > f19. The pistil and tori contained f1, f4, f6, f8,
embryos (18.9) > rhizomes (8.5) (Jung et al. f14, f15, f16, f17, f18 and f19. The flower stalks
2008). contained f1, f4, f6, f8, f10, f15, f18 and f19; the
Total flavonoid content was highest in the leaves contained f1, f4, f6, f11, f15, f16, f18 and
leaves 2.06 E3mg/100 g FW, followed by flower f20. Lotus seed pods contained f4, f6, f10, f11,
petals, stamens, plumule 402.9–496 mg/100 g f14, f15, f16, f17, f18 and f19; seed coats con-
FW and seed kernel the lowest 0.67 mg/100 g tained f2, f3, f4, f7, f9, f11, f12, f13, f14, f15, f16,
FW (Li et al. 2014). Thirty-three flavonoids were f17, f18 and f19. The seed kernels contained f9,
found in lotus flowers, seeds and leaves: f11, f12, f15, f16 and f18. Thirteen C-glycosyl
myricetin 3-O-galactoside (f1), luteolin 8-C-β-d- flavones (newly isolated C-glycosides, f7, f9,
glucopyranoside (orientin) (f2), luteolin 6-C-β-d- f21, f22, f23, f24, f25, f26, f27, f29 and f31) and
glucopyranoside (isoorientin) (f3), myricetin six O-glycosides (newly isolated O-glycosides,
3-O-glucuronide (f4), myricetin 3-O-glucoside f20, f28, f30, f32 and f33) within plumules were
(f5), quercetin 3-O-arabinopyranosyl-(1 → 2)- detected.
galactopyranoside (f6), apigenin 8-C-β-d- The extracts of lotus rhizomes, seeds, flowers
glucopyranoside (vitexin) (f7), quercetin 3-O-α- and leaves had been reported to have varied ther-
l-rhamnopyranosyl-(1 → 6)-β-d-glucopyranoside apeutic potential (Mukherjee et al. 2009). Several
(rutin) (f8), apigenin 6-C-β-d-glucopyranoside bioactive compounds had been derived from
(isovitexin) (f9), quercetin 3-O-β-d-galactopy- these plant parts belonging to different chemical
ranoside (hyperoside) (f10), quercetin 3-O- groups, including alkaloids, flavonoids, glyco-
glucuronide (f11), quercetin 3-O-β-d-glucopy- sides, triterpenoid, vitamins, etc., which all
ranoside (isoquercitrin) (f12), kaempferol 3-O- imparting their own therapeutic impact.
76 Nelumbonaceae
20 and 50 kGy, while the Hunter colour b*-val- petroleum extracts had comparable AAC with
ues were decreased under the same conditions. BHA by the beta-carotene bleaching assay.
Lotus leaf extract exhibited 2-diphenyl-1- Lotus rhizome knot extract exhibited high
picrylhydrazyl hydrate (DPPH) free radical antioxidative capacity, as measured by1-
scavenging activities increased in a concentra- diphenyl-2-picrylhydrazyl and 2,2′-azino
tion-dependent manner (Ha et al. 2010). Mouse (3-ethylbenzothioazolino-6-sulfonate) and mea-
embryonic fibroblast (MEF) cells, damaged by sured by electron spin resonance (ESR) trapping
oxidative stress, decreased their viability follow- of the transient carbon-centred 1-hydroxyethyl
ing increasing concentration of H2O2, but the radical (generated in a Fenton-type reaction) (Hu
cotreatment of n-butanol fraction of the leaf and Skibsted 2007). Whole rhizome, however,
extract and H2O2 resulted in an increase in cell only showed a significantly high scavenging
growth, by about 25 %, compared to the cells activity for small carbon-centred radicals, as
treated with H2O2. At 100 μg/ml extract concen- measured by the ESR method. Total phenol con-
tration, lotus leaf, flower and seed extracts tent in the plant extract correlated with the anti-
showed 78 %, 78 % and 80 % DPPH free radical oxidant capacity, except for the scavenging of
scavenging activity (Kim et al. 2011). The IC50 carbon-centred radical. Studies found that
values (μg/ml) of the antioxidation effects were gamma radiation treatment significantly reduced
flower, 5 μg; seed, 5 μg; leaf, 29 μg; root, 150 μg; microbial load, enhanced the DPPH scavenging
stem(rhizome), 162 μg; and vitamin C, 5 μg. activity and increased the storability of the irradi-
The amounts of phenolics, flavonoids and pro- ated lotus rhizome samples (Khattak et al. 2009).
anthocyanidins in the lotus leaf extracts from The treated samples were also acceptable sen-
various areas in China varied widely, ranging sorically. The extraction yield and phenolic con-
from 354 to 487 mg/g gallic acid equivalents, tents increased with the increase of radiation
from 172 to 236 mg/g rutin equivalents and from dose.
124 to 179 mg/g catechin equivalents, respec- The lotus rhizome showed the strongest anti-
tively (Huang et al. 2010a). All of the extracts oxidant activity in both assays of lipid peroxida-
had strong antioxidant activity in comparison to tion and inhibition of lysis of erythrocytes
the standard compounds butylated hydroxytolu- induced by peroxyl radicals (Jiang et al. 2010).
ene and vitamin C. Wild lotus samples from Of three fractions, L2 fraction showed the high-
Baiyangdian Lake and Weishan Lake exhibited a est antioxidant activity and was further fraction-
stronger free radical scavenging effect and greater ated, and L2c showed the strongest activity in
reducing power than the cultivated samples, but inhibiting haemolysis of erythrocytes, and on
no such differences were observed in the inhibi- further purification, L2c-3 was purified and
tion of lipid oxidation. identified as tryptophan. Its inhibitory concen-
tration of 50 % (IC50) value in inhibiting hae-
Lotus Rhizome molysis of erythrocytes was 156.3 μg/ml (i.e.
Methanol extract of lotus rhizome powder 765.4 μM).
showed the highest extract yield among all of sol-
vents (Yang et al. 2007). Although acetone Lotus Flower
extract had the highest total phenolics content, Ethanol lotus flower extract exhibited higher
methanol extract had the highest total phenolics antioxidant activity in-vitro (lower IC50 value)
recovery from lotus powder (20.1 mg catechin than vitamin C (Gayathri et al. 2009). A metha-
equivalents/100 g lotus powder). Extract of either nol extract of lotus stamens showed strong anti-
methanol or acetone demonstrated the highest oxidant activity in the peroxynitrite (ONOO-)
DPPH scavenging activity at both 66.7 and system and marginal activity in the DPPH and
133.3 mg/L. All extracts exhibited higher anti- total reactive oxygen species (ROS) systems
oxidant activity coefficient (AAC) than that of (Jung et al. 2003). After fractionation, the ethyl
ascorbic acid; furthermore, dichloromethane and acetate fraction exhibited strong antioxidant
Nelumbo nucifera 79
tive regulation on IL-2, IL-4 and IL-10. 3-O-glucuronide (Q3GA), a major quercetin
Furthermore, L2f-3 and LB2 might inhibit HIV-1 metabolite after absorption from the digestive
directly by downregulating TNFα (tumour necro- tract, enhanced ATP-binding cassette, subfam-
sis factor α). ily A and member 1 (ABCA1) expression, in-
vitro, via LXRα in macrophages. ABCA1 is a
crucial cholesterol transporter involved in
Antiobesity/Antihyperlipidaemic/ reverse cholesterol transport to produce high-
Anticholesterolemic Activities density lipoprotein-cholesterol.
sues in rats fed a high-fat diet (You et al. 2014b). that lotus leaf extract had potential as an antiobe-
Consumption of a high-fat diet resulted in an sity agent by inhibiting pancreatic lipase and adi-
increase in serum total cholesterol (TC) and tri- pocyte differentiation (Ahn et al. 2013). Among
glyceride (TG) levels; however, administration of its constituents, the alkaloids trans-N-
lotus extract resulted in a decrease in the levels of coumaroyltyramine and cis-N-feruloyltyramine
TC and TG. Administration of the extract also significantly inhibited pancreatic lipase, whereas
resulted in a decrease in the level of serum leptin alkaloids (6R, 6aR)-roemerine-Nβ-oxide and liri-
and insulin and in hepatic thiobarbituric acid odenine showed strong inhibitory effect on adi-
reactive substance content, elevated by a high-fat pocyte differentiation.
diet and caused an increase in superoxide dis-
mutase activity and hepatic glutathione content.
The results suggested that lotus root exerts anti- Antidiabetic Activity
oxidant and antiobesity effects and could be used
as a functional and nutraceutical ingredient in Lotus rhizome extract exhibited hypoglycaemic
combating obesity-related diseases. effect in streptozotocin-induced diabetic rats
Administration of Jiang-Zhi-Ning (a tradi- (Mukherjee et al. 1995c). The LD50 of the extract
tional Chinese medicine containing fleeceflower was found to be 2 g/kg. The extract (300 mg/kg
root, Fructus crataegi, Folium nelumbinis and and 600 mg/kg, orally) caused a reduction of
Semen cassiae) extract and effective fraction sig- blood glucose levels in streptozotocin-induced
nificantly reduced contents of serum total choles- diabetic rats by 53 % and 55 %, respectively, at
terol (TC), triglyceride (TG), low-density the end of 12 h.
lipoprotein-cholesterol (LDL-C), coronary index The crude protein of lotus seeds caused a
and atherogenic index in hyperlipidaemic rats as significant decrease (44.80 %) in the blood glu-
well as significantly increased contents of high- cose level of diabetic albino rats after 2 weeks
density lipoprotein-cholesterol (HDL-C), in rats of treatment (Ibrahim and El-Eraqy 1996). Oral
(Chen et al. 2011). Moreover, they significantly administration of lotus rhizome ethanolic
enhanced the activity of serum total nitric oxide extract significantly markedly reduced the
synthase (NOS) and increased contents of nitric blood sugar level of normal, glucose-fed hyper-
oxide. They also caused significant reductions in glycaemic and streptozotocin-induced diabetic
contents of endothelin-1 and malondialdehyde as rats, when compared with control animals
well as significant increase in superoxide dis- (Mukherjee et al. 1997b). The extract improved
mutase (SOD) activity and total antioxidant glucose tolerance and potentiated the action of
capacity (T-AOC) in the hyperlipidaemic rats. exogenously injected insulin in normal rats.
In-vitro, they restored and enhanced the vitality When compared with tolbutamide, the extract
of HUVECs with a concentration-dependent exhibited activity of 73 and 67 % of that of tol-
manner as well as content of NO in the culture butamide in normal and diabetic rats, respec-
media of HUVEC. They caused reductions in the tively. A methanol extract of the lotus stamens
contents of endothelin-1 and malondialdehyde in exerted an inhibitory effect on rat lens aldose
HUVECs. They also caused an increase in the reductase (RLAR) which had been shown to
vitality of SOD and T-AOC in HUVECs. play an important role in the complications asso-
Furthermore, they enhanced LDL-RmRNA ciated with diabetes (Lim et al. 2006). Among the
expression andCYP7A1 mRNA with in a isolated flavonoids, those harbouring 3-O-α-l-
concentration-dependent manner. Finally, they rhamnopyranosyl-(1 → 6)-β-d-glucopyranoside
caused reductions in the contents of cholesterol groups in their C rings, including kaempferol
in Bel-7402. They also increased intercellular 3 - O - α - l - r h a m n o py r a n o s y l - ( 1 → 6 ) - β - d -
content of total bile acid as well as lowered extra- glucopyranoside and isorhamnetin 3-O-α-l-
cellular contents of TBA in the cells in a rhamnopyranosyl-(1 → 6)-β-d-glucopyranoside,
concentration-dependent manner. Studies found exhibited the highest degree of rat lens aldose
84 Nelumbonaceae
reductase inhibitory activity in-vitro, with IC50 val- or symptoms of toxicity or mortality. The find-
ues of 5.6 and 9.0 μM, respectively. ings indicated that lotus flower extract had non-
Studies by Pan et al. (2009) demonstrated that acute toxicity but possessed hypoglycaemic and
neferine, from lotus embryo, had effects similar hypolipidaemic activities.
to rosiglitazone in significantly decreasing fast- Among several compounds extracted from
ing blood glucose, insulin, triglycerides and antidiabetic plants, nuciferine from N. nucifera
tumour necrosis factor-α (TNF-α) and enhancing was found to stimulate insulin secretion in iso-
insulin sensitivity in insulin-resistant rats. lated beta INS-1E cells by blocking potassium-
Streptozotocin-induced diabetic rats treated with adenosine triphosphate channels, thus
lotus seed ash at a concentration of 200 mg/kg contributing to the antidiabetic effects of
body weight orally for 30 days exhibited signifi- Nelumbo nucifera (Nguyen et al. 2012). Studies
cant hypoglycaemic activity (Mani et al. 2010). showed that lotus plumule polysaccharide (LPPS)
The hypoglycaemic activity of the ash was com- administration on non-obese diabetic (NOD)
parable with gliclazide. The presence of trace female mice for 15 weeks significantly increased
elements in appreciable amounts in the seeds pancreatic islet cell numbers and slightly
may play a direct or indirect role on insulin secre- enhanced the basal insulin secretion ability com-
tion or its action in a synergetic manner. Lotus pared to the control group (Liao and Lin 2013a).
leaf methanolic extract (NNE) increased insulin LPPS administration improved serum lipid pro-
secretion from β cells (HIT-T15) and human files in the diabetic mice via relatively increasing
islets in-vitro (Huang et al. 2011a). NNE serum high-density lipoprotein-cholesterol, but
enhanced the intracellular calcium levels in β decreasing low-density lipoprotein-cholesterol
cells and could also enhance phosphorylation of and total cholesterol levels. All four lotus leaf
extracellular signal-regulated protein kinases compounds, 2-hydroxy-1-methoxyaporphine,
(ERK)1/2 and protein kinase C (PKC). In-vivo pronuciferine, nuciferine and roemerine, showed
studies showed that NNE possessed the ability to the effects of improving insulin-stimulated glu-
regulate blood glucose levels in fasted normal cose consumption in differentiated 3T3-L1 adi-
mice and high-fat diet-induced diabetic mice. Of pocytes compared with the control group (Ma
its active constituents, quercetin did not affect et al. 2014). 2-hydroxy-1-methoxyaporphine and
insulin secretion, but catechin significantly and pronuciferine exhibited the most potent glucose
dose dependently enhanced insulin secretion. consumption-stimulatory activity at the concen-
Orally administered catechin significantly tration of 2 μg/mL.
reversed the glucose intolerance in high-fat diet-
induced diabetic mice. The findings suggested
that NNE and its active constituent catechin were Anti-inflammatory Activity
useful in the control of hyperglycaemia in non-
insulin-dependent diabetes mellitus through their Kaempferol 3-O-glucoside and luteolin
action as insulin secretagogues. Administration 7-O-glucoside from lotus aerial parts exhibited
of lotus flower extract at a dose of 250 mg/kg sig- anti-inflammatory activity (Wassel et al. 1996).
nificantly decreased the levels of fasting blood Methanol lotus rhizome extract at doses of 200
glucose (FBG), total cholesterol (TC), triglycer- and 400 mg/kg and betulinic acid at doses of car-
ides (TG), low-density lipoprotein (LDL) and rageenin and serotonin induced rat paw oedema
blood urea nitrogen (BUN) but increased high- in rats (Mukhjee et al. 1997a). The effects pro-
density lipoprotein (HDL) in streptozotocin- duced were comparable to that of phenylbuta-
induced diabetic rats compared to untreated zone and dexamethasone, two prototype
diabetic controls (Sakuljaitrong et al. 2013). anti-inflammatory drugs. Studies showed that
However, the extract did not alter creatinine in kaempferol, constituent of lotus extract, when
diabetic-treated rats compared to diabetic con- administered to rat tissues, attenuated the NF-κB
trols. Also the extract did not produce any signs nuclear binding activity (Kim et al. 2007).
Nelumbo nucifera 85
induced sleeping time. An anxiolytic, diazepam lotus embryos, inhibited mice locomotor activity
decreased locomotor activity and rectal tempera- indicating that they all possessed sedative activ-
ture and enhanced sleep elicited by thiopental, ity (Nishimura et al. 2013). The efficacy of lien-
similar to neferine. In addition, neferine and sinine, isoliensinine and nelumboferine was more
diazepam showed anti-anxiety effects in the ele- potent than neferine and O-methylneferine.
vated plus maze test. Neferine did not affect mus- Compounds isolated from lotus leaves
cle coordination by the rotarod test. Neferine did 6′-hydroxy-4,4′-dimethoxychalcone, rhamnetin-
not affect strychnine- nor picrotoxin-induced sei- 3-O-β-d-glucopyranoside, quercetin-3-O-β-d-
zure. The results suggested that neferine had sev- glucopyranoside, quercetin-3-O-α- l-
eral central nervous system effects and that rhamnopyranosyl, quercetin-3-O-rutinoside and
neferine may participate in the efficacy of the isorhamnetin-3- O -α- l - rhamnopyranosyl-
sedative effects of embryos of N. nucifera seeds. (1 → 6)-[α- d - lyxopyranosyl-(1 → 2)-β- d -
Results of studies showed that lotus seed extract glucopyranoside] showed inhibitory activities
ameliorated scopolamine-induced amnesia in against beta amyloid (1–42) by A-beta aggrega-
rats by inhibiting acetylcholinesterase activity tion method with inhibition rates of (63.99)%,
and inducing choline acetyltransferase expres- (79.61)%, (49.96)%, (101.19)%, (88.41)% and
sion (Oh et al. 2009). The i.p. administration of (72.48)%, respectively (Zhao et al. 2013b). Of
neferine 25–100 mg/kg elicited anti-immobility the alkyl 4-hydroxybenzoates isolated from lotus
effects in mice (Sugimoto et al. 2010). The seeds, methyl 4-hydroxybenzoate and ethyl
molecular dose effects of neferine in the forced 4-hydroxybenzoate enhanced 5-HT-stimulated
swimming test were almost equal to those of the inward current (I5-HT) in Xenopus oocytes, but butyl
typical antidepressants maprotiline and imipra- 4-hydroxybenzoate reduced it (Youn et al. 2010).
mine. The antidepressant effect was found to be The results indicated that butyl 4-hydroxybenzoate
mediated by serotonin1A (5-HT1A) receptor and was an inhibitor of the 5-HT3A receptors
was likely to be linked to serotonergic expressed in Xenopus oocytes.
neurotransmission. N-methylasimilobine, a potent acetylcholines-
Compounds cycloartenol, p-hydroxybenzoic terase (AChE) from N. nucifera leaves, inhibited
acid, vanilloloside and nuciferoside, isolated 50 % of AChE activity at the concentrations of
from lotus stamens, exhibited good and noncom- 1.5 μg/mL when the IC50 value of physostigmine
petitive inhibition against acetylcholinesterase standard was 0.013 μg/mL (Yang et al. 2012).
(AChE) with IC50 values of 11.89, 20.07, 4.55 The mode of AChE inhibition by N-methyla-
and 3.20 microM and Ki values of 15.71, 25.44, similobine was reversible and noncompetitive.
7.76 and 5.76 μM, respectively (Jung et al.
2010b). Cycloartenol, p-hydroxybenzoic acid
and nuciferoside also possessed butyrylcholines- Cognitive-Enhancing Activity
terase (BChE) inhibitory activities with IC50 val-
ues of 13.93, 62.29, 205.78 and 83.06 μM, The methanol extract of N. nucifera rhizome elic-
respectively. The selectivity index (SI) values of ited in significant improvements of memory
1, 2, 3 and 5, calculated from IC50 values of BChE functions and neurogenesis in the dentate gyrus
and AChE, were 1.2, 3.1, 45.7 and 26.0. However, of Wistar rats (Yang et al. 2008). In the passive
all isolated compounds lacked BACE1 inhibition avoidance test, the retention time of extracted-
up to 100 μM. The data suggested that N. nucifera treated rats was significantly longer than that of
stamen-derived compounds could potentially controls. In addition, cell proliferation and cell
exert their primary anti-Alzheimer effects as differentiation increased significantly in the den-
AChE inhibitors rather than BACE1 inhibitors. tate gyrus. The results suggested that N. nucifera
Neferine, liensinine, isoliensinine, nelum- rhizome extract may improve learning and mem-
boferine and O-methylneferine, isolated from ory by enhancing neurogenesis in the dentate
Nelumbo nucifera 87
gyrus of the hippocampus. Hydroalcohol lotus increased the latency time measured during the
extract exhibited a weak inhibition of acetylcho- passive avoidance task in scopolamine-induced
linesterase with IC50value of 185.55 μg/mL, com- cholinergic blockade and normal naïve mice.
pared to physostigmine (IC50 0.076 μg/mL) used Allantoin treatment (3 or 10 mg/kg, for 7 days)
as a standard (Mukherjee et al. 2007). also increased the expression levels of phosphor-
Neferine, from lotus embryo, exerted signifi- ylated phosphatidylinositide 3-kinase (PI3K),
cant improvement in cognitive impairment in phosphorylated protein kinase B (Akt) and phos-
scopolamine-induced amnesia animal models phorylated glycogen synthase kinase-3β
and moderate inhibitory activities in cholinester- (GSK-3β). Also, allantoin significantly increased
ases (ChEs)- and β-site APP cleaving enzyme 1 the neuronal cell proliferation of immature neu-
(BACE1) assays (Jung et al. 2010a). Additionally, rons in the hippocampal dentate gyrus region.
it exhibited notable scavenging activities The results suggested that allantoin may have
against 1,1-diphenyl-2-picrylhydrazyl (DPPH), therapeutic potential for the cognitive dysfunc-
2,2′-azino-bis(3-ethylbenzothiazoline-6-sulfonic tions observed in Alzheimer’s disease.
acid) diammonium salt (ABTS), NO (nitric
oxide) and O2− (superoxide) radicals, as well as
ONOO− (peroxynitrite). Neferine also demon- Convulsant Activity
strated dramatic inhibitory activity against lipid
peroxidation and protein nitration in cell-free Nornuciferine derivatives administered intraperi-
antioxidant assays and moderate inhibitory activ- toneally to mice as hydrobromide or hydrochlo-
ity of NO generation with exceptional suppres- ride provoked intense clonic convulsions
sion of NF-κB activation in cell-based assays. (Burkman and Cannon 1972). Unlike apomor-
The results demonstrated that the anti-amnesic phine, they did not induce emesis in dogs.
effect of neferine may be mediated via antioxi- N-Propyl nuciferine was the most effective con-
dant and anti-inflammatory capacities, as well as vulsant agent.
inhibition of ChEs and BACE1. The administra-
tion of lotus rhizome extract significantly amelio-
rated the scopolamine-induced reduction of Analgesic Activity
Ki67- and doublecortin-immunoreactive cells in
the hippocampal dentate gyrus of rats (Yoo et al. Kaempferol 3-O-glucoside and luteolin
2011). In addition, the administration of the 7-O-glucoside from lotus aerial parts exhibited
extract significantly restored the scopolamine- analgesic activity (Wassel et al. 1996). Methanol
induced reduction of brain-derived neurotrophic seed extract of red and white lotus, fed to albino
factor (BDNF) in hippocampal dentate gyrus rats, elicited considerable analgesic effect in the
homogenates. The results suggested that lotus acute pain (tail flick) model in a concentration-
rhizome extract could ameliorate the dependent fashion (Chakravarthi and Gopakumar
scopolamine-induced reductions of cell prolifer- 2009). The higher dose group of white lotus seed
ation, neuroblast differentiation and BDNF (600 mg/kg) exhibited more pronounced activity
levels. than other extracts.
Lotus plant extract was one of several plants
used in Indian Ayurvedic system of medicine for
improving cognitive function that showed IC50 Cosmeceutical/Antityrosinase
values <100 μg/ml for acetylcholinesterase Activity
inhibitory activity (Mathew and Subramanian
2014). Allantoin, an active compound in lotus, Lotus plant parts were found to have whitening
exerted memory-enhancing effects in mice (Ahn effect as measured by tyrosinase inhibition assay
et al. 2014). Subchronic administration of allan- and DOPA-oxidase inhibition assay, and anti-
toin (1, 3 or 10 mg/kg, for 7 days) significantly wrinkle effect as measured by elastase inhibition
88 Nelumbonaceae
assay (Kim et al. 2011). DOPA-oxidase inhibi- dose-dependent manners. These hepatoprotective
tion effect (whitening effect) of N. nucifera’s effects of lotus seed extract against CCl4 and
leaf, seed and flower extract was 59 %, 57 % and AFB1 might result from its potent antioxidative
50 %, respectively. N. nucifera’s leaf, seed and properties. In-vitro studies showed that neferine
flower extract showed 56 %, 49 % and 54 % elas- efficiently inhibited cultured hepatic stellate
tase inhibition (antiwrinkle effect) at 200 μg/ml, (HSC-T6) cell activation and induced apoptosis
while adenosine indicated 26 % inhibition. Lotus by increasing Bax and caspase 3 mRNAs expres-
plant parts were found to have whitening effect as sion via the mitochondrial pathway (Ding et al.
measured by tyrosinase inhibition assay and 2011).
DOPA-oxidase inhibition assay and antiwrinkle Administration of hydroalcoholic lotus seed
effect as measured by elastase inhibition assay extract to Wistar rats at 100 and 200 mg/kg body
(Kim et al. 2011). DOPA-oxidase inhibition weight for 4 days prior to carbon tetrachloride
effect (whitening effect) of N. nucifera’s leaf, treatment caused a significant dose-dependent
seed and flower extract was 59 %, 57 % and increase in the level of superoxide dismutase and
50 %, respectively. N. nucifera’s leaf, seed and catalase and a significant decrease in the level of
flower extract showed 56 %, 49 % and 54 % elas- thiobarbituric acid reactive substances (TBARS),
tase inhibition (antiwrinkle effect) at 200 μg/ml, when compared to CCl4-treated control in both
while adenosine indicated 26 % inhibition. Water liver and kidney (Rai et al. 2006). These changes
cream including Nelumbo nucifera’s root, leaf, observed at 100 mg/kg body weight treatment
flower and stem extract did not cause significant were comparable to those observed for standard
skin irritation. Methanolic extracts from the vitamin E at 50 mg/kg treatment. Armepavine, a
flower buds and leaves of sacred lotus were found lotus compound, exerted anti-hepatic fibrogenic
to show inhibitory effects on melanogenesis in effects in-vitro and in-vivo (Weng et al. 2009). It
theophylline-stimulated murine B16 melanoma dose dependently attenuated tumour necrosis
4A5 cells (Nakamura et al. 2013). Among the factor-α (TNF-α)- and lipopolysaccharide (LPS)-
constituents isolated, nuciferine, N-methyla- stimulated α-SMA protein expression and AP-1
similobine, (–)-lirinidine and 2-hydroxy-1-me- activation by HSC-T6 cells without adverse cyto-
thoxy-6a,7-dehydroaporphine showed potent toxicity in bile duct-ligated (BDL) rats. It signifi-
inhibition of melanogenesis. In addition, cantly reduced plasma AST and ALT levels,
3–30 μM nuciferine and N-methylasimilobine hepatic α-SMA expression and collagen contents
inhibited the expression of tyrosinase mRNA, and fibrosis scores as compared with vehicle
3–30 μM N-methylasimilobine inhibited the treatment. The results suggested that armepavine
expression of TRP-1 mRNA and 10–30 μM both in-vitro and in-vivo has antifibrotic effects
nuciferine inhibited the expression of TRP-2 in rats, possibly through anti-NF-κB activation
mRNA. pathways. Oral administration of lotus root hot
water extract with taurine supplementation
exerted antioxidant and hepatic protective effects
Hepatoprotective Activity in high-fat diet-induced obese male Sprague–
Dawley rats (Du et al. 2010). Activities of serum
Ethanol lotus seed extract showed potent DPPH glutamate oxaloacetate transaminase, glutamate
free radical scavenging effects with a median pyruvate transaminase and thiobarbituric acid
inhibition concentration of 6.49 g/ml (Sohn et al. reactive substance content were reduced, and
2003). Treatment of hepatocytes with the extract hepatic antioxidant enzymes were reduced by
inhibited both the production of serum enzymes lotus root extract. An ethanolic lotus leaf extract
and cytotoxicity by carbon tetrachloride (CCl4). at doses of 300 and 500 mg/kg exhibited hepato-
The genotoxic and cytotoxic effects of aflatoxin protective activity against CCl4-induced liver
B1 (AFB1) were also inhibited by the extract in toxicity in rats and antioxidant activity at 100 mg/
Nelumbo nucifera 89
kg that was comparable with that of a standard ameliorated high-fat diet-induced dyslipidaemia
treatment comprising 100 mg/kg of silymarin, a as well as liver steatosis and hepatic injury (Guo
known hepatoprotective drug (Huang et al. et al. 2013). The beneficial effects of nuciferine
2010b). were associated with altered expression of
Lotus germ oil treatment of mice with chronic hepatic genes involved in lipid metabolism. Tang
hepatotoxicity induced by carbon tetrachloride et al. (2014) showed that the polyphenol-rich
increased significantly the activities of antioxi- lotus leaf extract inhibited alcohol-induced ste-
dant enzymes, such as superoxide dismutase atohepatitis by multiple ways including reducing
(SOD), catalase (CAT) and the concentration of hepatic lipid synthesis, oxidative stress and anti-
glutathione (GSH) in a dose-dependent manner inflammation and enhancing fatty acid oxidation
(Lv et al. 2012). Also the lipid peroxidation prod- and transport responses in C57BL/6 J mice.
uct malondialdehyde and serum levels of hepatic
enzyme biomarkers (alanine aminotransferase
and aspartate aminotransferase) were decreased. Pulmonoprotective Activity
In addition, lotus germ oil could inhibit the con-
version of super-coiled pBR322 plasmid DNA to Administration of isoliensinine, a constituent of
the open circular form and apoptosis of hydrogen lotus embryo, markedly suppressed the increase
peroxide-induced PC-12 cells. in hydroxyproline content and abated pulmonary
After 3 weeks of feeding, the hepatomegaly fibrosis histological injury induced by bleomycin
and hepatic triglyceride accumulation was mark- in mice (Xiao et al. 2005). Isoliensinine enhanced
edly alleviated in the lotus polyphenol-diet-fed serum superoxide dismutase activity and
db/db mice relative to the control mice (Tsuruta decreased the bleomycin-elevated malondialde-
et al. 2011, 2012). Although the lipolytic enzyme hyde level in a concentration-dependent manner.
activity was not changed, the activities of lipo- Moreover, isoliensinine also significantly inhib-
genic enzymes, such as fatty acid synthase and ited the overexpression of TNF-α and TGF-β1
malic enzyme, were significantly lower in the induced by bleomycin. The results indicated that
lotus polyphenol-diet-fed db/db mice. Further, isoliensinine possessed a significant inhibitory
the presence of B-type proanthocyanidin poly- effect on bleomycin-induced pulmonary fibrosis,
mers with polymerisation degree up to 9 was probably due to its antioxidant and/or anti-
detected in the polyphenolic lotus root extract. It inflammatory activities and its inhibition of over-
was suggested that condensed tannins contained expression of TNF-α and TGF-β1 induced by
in lotus root could alleviate hepatic steatosis by bleomycin. Neferine, from lotus embryo, signifi-
suppressing the lipogenic enzyme activity in the cantly inhibited amiodarone-induced pulmonary
livers of db/db mice. They also found that the fibrosis in adult Kunming mice modulated by its
serum levels of adiponectin, which had been properties of anti-inflammation, surfactant pro-
reported to have a protective effect against non- tein-D inhibition and restoring increased
alcoholic fatty liver disease (NAFLD), were sig- CD4+CD25+ regulatory T cells which may regu-
nificantly higher in the Lotus group than in the late Th1/Th2 imbalance by suppressing Th2
control group of the db/db mice. Additionally, the response (Niu et al. 2013).
hepatic expression of such inflammatory genes as
tumour necrosis factor-alpha and monocyte che-
moattractant protein-1 was markedly suppressed Renoprotective Activity
by the Lotus diet. They speculated that the devel-
opment and progression of NAFLD were pre- Studies showed that (S)-armepavine, from lotus,
vented by suppressing the expression of lipogenic administered in the established phase of autoim-
and inflammatory genes as a result of the higher mune crescentic glomerulonephritis (ACGN) in
serum adiponectin level in the Lotus diet-fed db/ mice, was capable of markedly decreasing glo-
db mice. Nuciferine supplementation in hamsters merular crescents in the kidney and improving
90 Nelumbonaceae
proteinuria and renal dysfunction (Ka et al. action; its mechanism may be related to blockade
2010). These therapeutic effects were associated of Ca2+, Na+ influx (Wang et al. 1992). Liensinine
with (1) early systemic negative modulation of 3 mg/kg i.v. temporarily inhibited all parameters
T/B cells; (2) intra-renal regulation of combined of haemodynamics in anesthetised or pithed rats.
NF-κB activation and mononuclear leukocytic The dose-dependent inhibitory effects of liensi-
infiltration, thereby preventing glomerular cres- nine (1–30 mg/kg) on LVP, +dp/dtmax and SAP
cent formation; and (3) protection from apoptosis (slow action potential) in anesthetised rats were
in both the spleen and kidney. slightly stronger than those of quinidine 3 mg/kg.
Liensinine 1–30 mg/kg dose dependently pro-
duced these actions. The magnitude of inhibi-
Spasmolytic/Anti-arrhythmic/ tory effect of liensinine on all haemodynamic
Hypotensive/Bradycardic Activities parameters nearly corresponded to these of
verapamil 1 mg/kg. Liensinine 1–100 µmol/L
Nn-9 a non-crystalline alkaloid isolated from reduced the contractile force of isolated left
lotus embryo exerted hypotensive effects in ani- atria and the spontaneously beating rate of iso-
mals (Chen et al. 1962). In anesthetised cats, i.v. lated right atria of rabbits in concentration-
injections of 1–2 mg/kg lowered, after a delay of dependent manner. Liensinine 10–100 µmol/L
1/2–1 min, the blood pressure by about 50 % was also shown to concentration dependently
from the original level, lasting 2–3 h. In dogs, the decrease the action potential amplitude (APA)
blood pressure returned to normal in 1/2 h. This and the maximal velocity of phase 0 depolarisa-
dosage was ineffective in rabbits. Pretreatment tion (Vmax) and prolong the sinus cycle length
with Nn-9 also lessened the hypotensive action of (SCL) of SAP in isolated sinoatrial node pace-
histamine. Nn-9 appeared to act primarily maker cells of rabbits (Wang et al. 1993).
through histamine release. The alkaloids asimilo- Moreover, liensinine 1–100 µmol/L was found
bine and lirinidine, isolated from lotus leaves, to inhibit the slow inward current of canine car-
inhibited the contraction of isolated rabbit aorta diac Purkinje fibre in concentration-dependent
induced by serotonin; the pA2 values were 5.78 manner. Liensinine 3 and 100 µmol/L reduced
and 7.36, respectively (Shoji et al. 1987). the peak value of slow inward current by 14 %
Studies found that neferine from lotus embryo and 88 %, respectively. The results suggested
possessed antiarrhythmic action (Li et al. 1989). that liensinine possessed calcium antagonistic
It exerted an inhibitory effect on the slow trans- effects.
membrane Na+ and/or Ca2+ current of myocar- Studies by Morales et al. (1997) demonstrated
dium. Neferine decreased the amplitude of action the hypotensive and bradycardic properties of
potential and the maximal upstroke velocity in (±)-norarmepavine, a lotus alkaloid. In rat’s aorta,
rabbit sinoatrial nodes and in the clusters of cul- (±)-norarmepavine (10 mg/kg i.v.) decreased the
tured cardiac myocytes from neonatal rats in a mean arterial pressure and heart rate by 45 % and
concentration-dependent manner. In subsequent 21 %, respectively. It showed a negative chrono-
studies, neferine at 1–10 mg/kg i.v. dose depend- tropic effect on rat-isolated atria, decreasing the
ently decreased the monophasic action potential spontaneous frequency by about 54 %. Aortic
amplitude, prolonging the monophasic action rings contracted with KCl 70 mM were relaxed
potential duration in anaesthetised cat’s heart (Li in a concentration-dependent manner by
et al. 1990). It also decreased left ventricular (±)-norarmepavine, (±)-coclaurine and (±)-norco-
pressure (LVP), dP/dt, prolonged sinus cycle claurine. The two earlier alkaloids exhibited an
length and reduced arterial blood pressure in a efficacy similar to verapamil, relaxing the aortic
dose-dependent manner. These effects were simi- rings by 100 %. (±)-Norcoclaurine exhibited a
lar to those of quinidine and different from tet- lower efficacy. The rank order of potency was
randrine. Liensinine, an alkaloid from lotus verapamil > (±)-norarmepavine > (±)-norcoclau-
embryo, was shown to have antiarrhythmic rine > (±)-coclaurine.
Nelumbo nucifera 91
Neferine alkaloid isolated from lotus embryo aggregation is related to regulation of TXA2/
exhibited antiarrhythmic effect in experimental PGI2 and cAMP/cGMP balance. Studies found
arrhythmic models (Qian 2002). In guinea pig neferine could dose dependently inhibit human
papillary muscles and atria, neferine reduced the or rat platelet aggregation induced by ADP, col-
force of contraction, decreased the amplitude and lagen and epinephrine both in-vitro and in-vivo
upstroke velocity Vmax of action potential and (Qian 2002). Neferine, from lotus embryo, sig-
prolonged action potential duration (APD50 and nificantly and dose dependently inhibited colla-
APD 90) and effective refractory period. Neferine gen-, thrombin- and U46619-induced platelet
also suppressed the contractility of rabbit papil- aggregation in mice-washed platelets, or ADP-
lary muscles, prolonged functional refractory induced platelet aggregation in mice platelet-rich
period and inhibited automaticity, positive stair- plasma (Zhou et al. 2013b). Neferine treatment
case and post-rest potentiation. Neferine, isolated decreased platelet dense granule secretion initi-
from the seed embryo, increased the basal cAMP ated by collagen, thrombin and U46619. Also,
concentration in rabbit corpus cavernosum tissue Neferine markedly and dose dependently pro-
in a dose-dependent manner (Chen et al. 2008). moted the dissociation of platelet aggregates pre-
The accumulation of cAMP induced by prosta- formed by various agonists including collagen,
glandin E1 (PGE1, a stimulator of cAMP produc- thrombin, U46619 or ADP. Neferine could sig-
tion) was also augmented by neferine in a nificantly reduce the area of mice platelet adhe-
dose-dependent manner. It was found that nefer- sion to the collagen and inhibited thrombosis
ine with its spasmolytic activity could enhance in-vitro. In collagen–epinephrine-induced acute
the concentration of cAMP in rabbit corpus cav- pulmonary thrombus mouse model, neferine, at
ernosum tissue, probably by inhibiting phospho- 6 mg/kg, significantly attenuated thrombus
diesterase activity. Neferine, from lotus embryo, formation.
induced a concentration-dependent decrease in
current amplitude according to the voltage steps
and tail currents of human ether-à-go-go-related Immunomodulatory Activity
gene (HERG) K+ channels in a dose-dependent
manner with an IC50 of 7.419 μM (Gu et al. 2009) N. nucifera ethanolic extracts significantly sup-
in human embryonic kidney (HEK293) cells. It pressed phytohemagglutinin-activated human
had no effect on the generation and trafficking of peripheral blood mononuclear cell (PBMC) pro-
HERG protein. A blocked-off HERG channel liferation (Liu et al. 2004). The inhibitory action
was one mechanism of the antiarrhythmic effects of lotus extract did not involve direct cytotoxic-
by neferine. ity. The suppressant effects of lotus extract on
proliferation of activated PBMC appeared to be
mediated, at least in part, through inhibition of
Antithrombotic/Antiplatelet Activity early transcripts of PBMC, especially those of
important interleukin IL-2, interferon-gamma
Neferine was shown to significantly inhibit rabbit (IFN-gamma) and cyclin-dependent kinase (cdk)
platelet aggregation induced by ADP, collagen, 4 mRNA expression and arrest of cell cycle pro-
arachidonic acid (AA) and platelet-activating gression in the cells. Oral administration of MRL/
factor (PAF) with IC50 of 16, 22, 193 and MpJ-lpr/lpr mice with similar disease features to
103 μmol/L, respectively (Yu and Hu 1997). human systemic lupus erythematosus with
Neferine was found to increase vascular 6-keto- (S)-armepavine (from lotus) for 6 weeks pre-
PGF1 alpha and platelet cAMP levels in dose- vented lymphadenopathy and elongated life span
dependent manner, but inhibit AA-stimulated of MRL/MpJ-lpr/lpr mice (Liu et al. 2006). It
TXA2 release from platelets. The results sug- appeared to be mediated by inhibition of spleno-
gested that the mechanism of neferine on platelet cyte proliferation, suppression of interleukin-2
92 Nelumbonaceae
suggested that lotus leaf could be considered to temperature (pyrexia) in rats (Mukherjee et al.
have therapeutic value in the prevention of 1996a). The effect produced was comparable
atherosclerosis. with the standard antipyretic drug, paracetamol
(150 mg/kg, i.p.). The ethanol lotus stalk extract
at a dose of 200 mg/kg was found to produce sig-
Aldose Reductase Inhibitory Activity nificant lowering of normal body temperature up
to 3 h, and at 400 mg/kg, it caused significant
Among several solvent fractions of lotus leaf lowering of body temperature up to 6 h after its
extract, the ethyl acetate and n-butanol fractions, administration (Sinha et al. 2000). In the model
having prominent total phenolic content (TPC) of yeast-induced elevation of body temperature,
and total flavonoid content (TFC)C values, the extract showed dose-dependent lowering of
showed significant antioxidant effects in the body temperature up to 4 h at both the doses, and
DPPH and TEAC assays (Jung et al. 2008). the results were comparable to that of paracetamol,
Moreover, the ethyl acetate fraction exhibited a standard antipyretic agent.
superior inhibitory effects in the total reactive
oxygen species (ROS), rat lens aldose reductase
(RLAR) and advanced glycation end products Antihemorrhagic Activity
(AGE) assays, with IC50 values of 9.4, 2.4 and
28.2 μg/ml, respectively. Also, two of its key The antihemorrhagic principle in nelumbins
antioxidant flavonoids, quercetin 3-O-β-d- receptaculum, dried receptacle of Nelumbo
glucopyranoside and quercetin 3-O-β-d- nucifera, was identified as quercetin [2-(3, 4-dihy-
glucuronopyranoside, may play important roles droxy)-3, 5, 7-trihydroxy-4H-1-benzopyran-4-
in the antioxidant and RLAR inhibitory effects of one] (Ishida et al. 1988).
N. nucifera leaves. The hydroalcoholic extract of
lotus plant exhibited aldose reductase inhibitory
activity with an IC50 value 59.78 μg/mL, and its Antiallergic Activity
alkaloidal extract was more inhibitory with an
IC50 of 28.82 μg/mL (Gupta et al. 2014). Flow cytometric analysis revealed that Fc epsilon
RI expression on human basophilic KU812F
human basophilic KU812F cell surface was sup-
Diuretic Activity pressed in a concentration-dependent manner
when the cells were cultured with kaempferol
The methanol extract of lotus rhizomes induced extracted from a methanolic extract of flavonoid-
significant diuresis in rats (Mukherjee et al. rich lotus stamens (Shim et al. 2009). Moreover,
1996b). Dose-dependent effects were observed in RTPCR analysis showed that the mRNA levels for
urine volume and electrolyte excretion. There was Fc epsilon RI alpha- and gamma-chains were
a significant increase in natriuretic and chlorouretic reduced as the result of kaempferol treatment in a
activity, but kaliuresis was less than natriuresis. concentration-dependent manner. Kaempferol
showed its suppressive effects on intracellular Ca2+
concentration and histamine release from anti-Fc
Antipyretic Activity epsilon RI alpha-chain antibody-stimulated cells
in a concentration-dependent manner. These
The methanolic lotus rhizome extract, in doses of observations indicated that kaempferol may exert
200, 300 or 400 mg/kg (p.o.), produced signifi- antiallergic effect via downregulation of Fc epsi-
cant dose-dependent lowering of normal body lon RI expression and degranulation.
temperature and yeast-induced elevation of body
94 Nelumbonaceae
Antiestrogenic/Antisteroidogenic doses (100, 200, 400 and 600 mg/kg body wt.)
Activity reduced not only the frequency of defecation,
wetness of faecal dropping and PGE2-induced
The petroleum ether lotus seed extract adminis- enteropooling but also the propulsive movements
tered to mice exhibited antiestrogenic, antipro- of charcoal meal significantly in rats (Talukder
gestational and contraceptive activity at a dose of and Nessa 1998). Aqueous lotus fruit extract was
3 mg/kg body weight (Mazumder et al. 1992). found to have strong significant anti-rotaviral
Oral administration of lotus seed petroleum ether activity with a 50 % inhibitory concentration
extract to sexually immature female rats and (IC50) <300 μg/ml (Knipping et al. 2012). A com-
mature male rats on alternate day for 15 days bination of Glycyrrhiza glabra and lotus showed
caused a remarkable delay in sexual maturation synergy in their antiviral activities and interven-
in prepubertal female rats as evidenced from age tions of lotus, and combination may be useful in
of vaginal opening and first estrus (cornified the treatment of diarrhoea caused by rotavirus
smear) and a significant reduction in the sperm infection.
count and motility in mature male rats (Gupta
et al. 1996). In both the cases, treatment resulted
in accumulation of cholesterol and ascorbic acid Anti-dermatitis Activity
and reduction in D5-3b-hydroxysteroid dehydro-
genase and glucose-6-phosphate dehydrogenase Oral administration of lotus leaf extract to
activity in the ovary and testis of female and male NC/Nga mice with atopic dermatitis induced
rat, respectively. The results indicated suppres- by repeated epicutaneous application of
sion of steroidogenesis in both ovary and testis by 2,4-dinitrochlorobenzene (DNCB) on the dorsal
lotus seed extract. skin resulted in the suppression of clinical sever-
Oral administration of lotus seed ethanol ity score, transepidermal water loss, scratching
extract to female albino rats elicited in significant behaviour and blood IgE level (Karki et al. 2012).
decline in weights of reproductive organs (ovary, Histopathologic analyses revealed that thicken-
uterus, vagina) (Mutreja et al. 2008). Glycogen ing of the epidermis and mast cell degranulation
levels in the reproductive organs were decreased were significantly reduced in lotus-treated mice.
significantly, but the cholesterol levels were The results suggested that lotus leaf extract may
increased; no changes in haematological param- be a useful natural resource for the management
eters were found. The extract prolonged the oes- of atopic dermatitis.
trous cycle. The results suggested that lotus seed
exhibited antiestrogenic activity without altering
the general physiology of the female rats. Antimalarial Activity
Prebiotic Activity 1.10 ppm; LC90 = 28.13, 35.83 and 3.59 ppm),
respectively. Of nine plant extracts tested ,the
Studies showed that prebiotics such as lotus seed- highest larvicidal activity was observed after
resistant starch, especially P-LRS3, could stimu- 24 h in the leaf methanol extract of N. nucifera,
late the growth of Bifidobacterium adolescentis seed ethyl acetate and methanol extract of P.
(Zhang et al. 2013a). Compared with glucose and nigrum against the fourth instar larvae of
high amylose maize starch (HAMS) media, bifi- Anopheles stephensi (LC50 = 34.76, 24.54 and
dobacteria had a higher tolerance to gastrointesti- 30.20 ppm) and against Culex quinquefasciatus
nal conditions in lotus-resistant starches LRS3 (LC50 = 37.49, 43.94 and 57.39 ppm), respec-
and P-LRS3 media. The rough surface of lotus- tively (Kamaraj et al. 2011). Lotus leaf ethyl
resistant starch and the short chain fatty acids acetate extract caused 100 % mortality of A. ste-
produced during fermentation might influence phensi and C. quinquefasciatus after 48 h expo-
the proliferation of bifidobacteria. Lotus seed- sure. The maximum repellent activity was
resistant starch was more effective than either observed at 500 ppm in methanol extracts of N.
glucose or HAMS in promoting the proliferation nucifera, ethyl acetate and methanol extract of P.
of bifidobacteria (Zhang et al. 2014). nigrum and methanol extract of Trachyspermum
ammi. Lotus leaf extracts were found to have
insecticidal (adulticidal) activity (Santhoshkumar
Anthelmintic Activity et al. 2011). In adults of Haemaphysalis bispi-
nosa, parasitic mortality of 80, 74, 72 and 100 %
Aporphine alkaloids from lotus leaves, liriode- was observed in lotus crude leaf hexane, ethyl
nine, lysicamine, (–)-anonaine, (–)-asimilobine, acetate, acetone and methanol extracts, respec-
(–)-cadaverine, (–)-N-methylasimilobine, tively. Maximum efficacy was observed in lotus
(–)-nuciferine, (–)-nornuciferine, (–)-roemerine, leaf methanol extract against the adults of the
7-hydroxydehydronuciferine and cepharadione B Acarina H. bispinosa and the hematophagous fly
killed Hymenolepis nana or reduced their sponta- Hippobosca maculata with LC50 and LC90 values
neous movements (oscillation/peristalsis) but had of 437.14 and 200.81 ppm, respectively.
no larvicidal effect or ability to halt spontaneous
parasite movement for 72 h of Anisakis simplex
(Lin et al. 2014). The above compounds showed Drug Interaction/Cytochrome
greater lethal anthelmintic efficacy on H. nana Inhibitory Activity
than against A. simplex.
Studies showed that nuciferine inhibited cyto-
chrome P450 1A2 (CYP1A2) activity in-vitro
Insecticidal/Larvicidal Activity and caused changes in the pharmacokinetic
parameters of phenacetin after oral administra-
The hexane, chloroform, ethyl acetate, acetone, tion to Wistar rats (Hu et al. 2010). Studies
methanol and aqueous lotus leaf extracts and syn- revealed that the alcoholic lotus leaf extract and
thesised silver nanoparticles using aqueous lotus its alkaloid fraction strongly inhibited human
leaf extract showed moderate larvicidal effects cytochrome CYP2D6 with an IC50 value of
against fourth instar larvae of Anopheles subpic- 12.05 μg/mL and 0.966 μg/mL, respectively, and
tus and Culex quinquefasciatus (Santhoshkumar weakly inhibited other isoenzymes (CYP2C9,
et al. 2011). Maximum efficacy was observed in CYP2C19, CYP2E1 and CYP3A4) (Ye et al.
crude methanol, aqueous and synthesised silver 2014a). The flavonoid fraction was found to
nanoparticles against the larvae of A. subpictus weakly inhibit CYP2D6. The three aporphine
(LC50 = 8.89, 11.82 and 0.69 ppm; LC90 = 28.65, alkaloids isolated from the alkaloid fraction,
36.06 and 2.15 ppm) and against the larvae of C. nuciferine, N-nornuciferine and 2-hydroxy-1-
quinquefasciatus (LC50 = 9.51, 13.65 and methoxyaporphine significantly inhibited
Nelumbo nucifera 97
CYP2D6 with IC50 values of 3.78, 3.76 and uronic acid and 2-hydroxy-1-methoxyaporphine-
3.15 μM, respectively. 2-O-sulphate. Following a single oral and
intravenous administration of total bisbenzyliso-
quinoline alkaloids from lotus seed embryo in
Mutagenic/Antimutagenic Activity rats, liensinine, isoliensinine and neferine were
detected in rat plasma (Huang et al. 2011b). The
The nitrite-treated methanol extract of Nelumbo lower limit of quantification can reach 0.03 μg/ml
nucifera flowers exhibited the high mutagenicity for the three compounds. Studies found that
on Salmonella typhimurium TA 98 and TA 100 neferine had no effect on amiodarone plasma
strains (Wongwattanasathien et al. 2010). All pharmacokinetics when it was co-administered
dichloromethane extracts of flowers including intragastrically with amiodarone, an antiarrhyth-
lotus decreased the mutagenicity induced by the mic agent (Wan et al. 2011). Studies in Caco-2
product of 1-aminopyrene nitrite model on both cells demonstrated that liensinine, neferine and
tester strains. isoliensinine, from lotus embryo, were substrates
of P-glycoprotein, whereas MRP2 cell was not
involved in the transport process, suggesting that
Toxicity Studies P-glycoprotein might be responsible for the
absorption and distribution of the 3 alkaloids (Yu
None of the lotus leaf alkaloid compounds were et al. 2013b).
cytotoxic to Vero cells up to a concentration of After administration of neferine to rats, one-
23.8 μg/mL (Agnihotri et al. 2008). The results compartmental pharmacokinetic parameters indi-
from both acute and subchronic oral toxicity cated that nuciferine had rapid distribution,
studies suggested that the oral lethal dose of lotus extensive tissue uptake and poor absorption into
stamen extract for male and female rats was in systemic circulation (Gu et al. 2014). The values
excess of 5000 mg/kg, and the no-observed- of absolute bioavailability were (3.8)%, (4.2)%
adverse-effect level (NOAEL) of the extract for and (3.9)% after oral administration of 2.0, 5.0
both male and female rats was considered to be and 10.0 mg/kg nuciferine and intravenous
200 mg/kg/day (Kunanusorn et al. 2011). administration of 0.2 mg/kg nuciferine in rats.
The results suggested that nuciferine was distrib-
uted into the brain, liver and adipose tissue after
Pharmacokinetic Studies intravenous administration. A total of 37 in-vivo
components were identified in rats after oral
Nornuciferine was found to be metabolised by rat administration of a lotus leaf flavonoid extract,
and rabbit liver microsomes to lysicamine (Smith including quercetin-3-O-glucuronide, quercetin-
and Sood 1971). Rat, rabbit and guinea pig liver 3-O-glucoside, quercetin-3-O-galacoside, quer-
microsomes also dealkylated N-alkylated analog cetin and kaempferol, as well as their methylation,
of nornuciferine. 2-Hydroxy-1-methoxyaporphin glucuronidation and sulfonation metabolites (Ye
(2H1M), an alkaloid from Nelumbo nucifera, one et al. 2014b).
of the herbal component of the Chinese
Traditional Medicine Jiang-Zhi-Ning (JZN), was
identified as the constituent showing the major
pharmacodynamic effect (Chen et al. 2011, Traditional Medicinal Uses
2012a). The major metabolites of 2H1M after
intragastric administration of JZN to beagle dogs Nelumbo nucifera is a medical plant used in tra-
were analysed and identified as N-demethyl-2- ditional Chinese medicine, and every part is uti-
hydroxy-1-methoxyaporphine-2-O-glycuronic lised and officially listed in the Chinese
acid, 2-hydroxy-1-methoxyaporphine-2-O-glyc- Pharmacopoeia as:
98 Nelumbonaceae
– Lianzi, Semen Nelumbinis, the dried mature (Nelumbo nucifera Gaertn) roots have been used
seeds used as a sedative and tonic. as an antidiabetic and an antiproliferative remedy
– Lianzixin, Plumula Nelumbinis, the dried (Park et al. 2009).
embryos from the mature seeds used as a sed- In folk medicines, seeds are used in the treat-
ative and haemostatic. ment of tissue inflammation, cancer, skin dis-
– Lianfang, Receptaculum Nelumbinis, the eases and leprosy; used as poison antidote,
dried receptacles used as a haemostatic. antiemetic and cooling agent; and generally pre-
– Lianxu, Stamen Nelumbinis, the dry stamens scribed to children as diuretic and refrigerant
collected when the lotus flowers bloom used (Chopra et al. 1986; Liu et al. 2004). The fruits
as an adstringent. and seeds of lotus are astringent and used to treat
– Heye, Folium Nelumbinis, the dry leaves used hyperdipsia, dermatopathy, halitosis, menorrha-
as a haemostatic. gia, leprosy and fever (Nadkarni and Nadkarni
– Oujie, Nodus Nelumbinis rhizomatis, the dry 1982). Seed powder mixed with honey is useful
nodes of lotus rootstock used as a haemostatic in treating cough, while roots with ghee (melted
(Tang and Eisenbrand 1992). After charring fresh butter), milk and gold promote strength,
the receptacles, leaves and rootstocks are also virility and intellect. Its rhizomes are used for
used against bleeding. According to Chinese treating pharyngopathy, pectoralgia, spermator-
traditional knowledge, medicinal uses of dif- rhoea, leucoderma, small pox, diarrhoea, dysen-
ferent part of lotus plants are common in the tery and cough (Mukherjee et al. 1997a, b; Ou
treatment of diarrhoea, tissue inflammation 1989). The stem is used in indigenous Ayurvedic
and haemostasis (Yu and Hu 1997). medicines as a diuretic and anthelmintic and to
treat strangury, vomiting, leprosy, skin diseases
Nelumbo nucifera leaf has been used for sum- and nervous exhaustion. Young leaves with sugar
mer heat syndrome as home remedy in Japan and are useful to treat rectal prolapse, and the leaves
China, and it has recently been used to treat obe- boiled with Mimosa pudica in goat’s milk can be
sity in China (Ono et al. 2006). Embryo of lotus used to treat diarrhoea. Nelumbo nucifera has
seeds is used in traditional Chinese drug called been used for the treatment of several disorders
‘Lian Zi Xin’, which primarily helps to overcome including skin disease, cough, inflammation,
nervous disorders, insomnia, high favours (with fever and many other disorders (Mukherjee et al.
restlessness) and cardiovascular diseases (e.g. 2010b; Ou 1989). The leaves are used as an effec-
hypertension, arrhythmia) (Chen et al. 2007). tive drug for haematemesis, epistaxis, haemoptysis,
Jiang-Zhi-Ning (JZN), a TCM, is composed of haematuria, metrorrhagia, fever and inflamma-
four Chinese herbs, i.e. fleeceflower root, Fructus tory skin conditions (Ou 1989).
Crataegi, Folium Nelumbinis and Semen Cassiae Nelumbo nucifera seed has been used as an
is used to strengthen blood circulation of coro- antiobesity agent in traditional Chinese herbal
nary artery, arrhythmia and hyperlipidaemia medicine (Pan et al. 2009). Hyperlipidaemia in
(Chen et al. 2011). Lotus leaf is known to be rodents can be treated with lotus leaves (La Cour
effective for ‘overcoming body heat’ and stop- et al. 1995; Onishi et al. 1984). Leaves also pos-
ping bleeding and is commonly used as a tradi- sess diuretic and astringent properties and help to
tional curing plant for the treatment of treat fever, sweating and strangury and as styptic
haematemesis, epistaxis, haemoptysis, haematu- (Chinese Materia Medica 1977).
ria and metrorrhagia in traditional Chinese medi- The Chinese administer the rhizomes for diar-
cine (Lee and Lee 2011). The herb of lotus rhoea and dysentery; the Cambodians make a tea
(Nelumbo nucifera) leaves is the traditional for menorrhagia (Burkill 1966). The Chinese
Chinese medicine He Ye, which is commonly used the embryo to reduce high fever and in the
used to treat sunstroke, assuage thirst and cure treatment of cholera, haemoptysis and spermator-
both diarrhoea and fever in China (Ye et al. rhoea. In Malayan medicine, it is highly esteemed
2014a, b). In traditional Korean medicine, lotus as a tonic and taken as tea for fever. The Malays
Nelumbo nucifera 99
used pounded petals for syphilis. They are astrin- vested 4–5 months after planting the lotus.
gent and taken for diarrhoea and vomiting in Additionally, lotus shoots provide substrate for
Java. The Chinese used them in facial cosmetic. growth of epiphytic algae that are consumed by
In India, they are used for fever and in the tilapia fish. Lotus rhizomes knot, as a waste from
Philippines for dysentery. The stamens are astrin- food production, will be a potential material for
gent and diuretic and used by the Chinese. They extracting antioxidants (Hu and Skibsted 2007).
are also used in cosmetics. In India, they are In India, the fibres from the stem are used as lamp
employed as an astringent and cooling medicine. wicks (Burkill 1966). A Thai cigarette may be
In Indonesia and India, the juice from the petioles wrapped in lotus petal to impart a dainty
and peduncles is used for treating diarrhoea. appearance.
In Vietnam, the seeds are used to treat dysen- Studies by Thongtha et al. (2014) found that
tery, spermatorrhoea, leucorrhoea, palpitation, phosphorus in domestic wastewater was removed
insomnia, general debility and anorexia (NIMM from the initial concentration of 1.038 mg/L to
1999). The plumules are useful in the therapy of 0.094 and 0.048 mg/L by N. nucifera and Cyperus
permanent mild fever, insomnia, thirst and hae- alternifolius, respectively, within 5 days.
moptysis. The leaves are beneficial in treating Domestic wastewater is a source of phosphorus
bloody stools, haematuria, gingival and subcuta- contamination that causes eutrophication when it
neous haemorrhage. The stamens are effective contaminates aquatic environments.
for metrorrhagia, haemoptysis, spermotorrhoea Lotus leaf extract inhibited algal bloom more
and insomnia. The achnes in decoction are used effectively than the stem (rhizome) extract (He
for treating dysentery and aphonia. The recepta- et al. 2013). At 25 g/L, the leaf extract exhibited
cle in decoction is useful as haemostatic in treat- highest inhibition rate of 71.33 % and 88.14 %
ing bloody stools, urination, leucorrhoea and against the algae Microcystis aeruginosa and
hypertension. Rhizome decoction is employed as Scenedesmus quadricauda, respectively, while
haemostatic in treating bloody stools, haematu- the inhibition rate for the rhizome extract was
ria, haematemesis, epistaxis and metrorrhagia. 49.78 % and 52.14 %, respectively.
NIMM (1999) also reported that in India, a nutri-
tious flour made from the rhizome is given to
children affected by diarrhoea, dysentery and Comments
dyspepsia. A paste made of powdered rhizome is
applied for tinea and other dermatoses. Lotus is revered as a sacred religious flower in
Hinduism and Buddhism. Lotus flower is the
national flower of India and is legendary in
Other Uses folklore and religious mythology. In Japan,
Moriyama City’s prefectural flower is the lotus.
The lotus is a very beautiful and attractive plant The lotus flower is the national flower of Vietnam
cultivated in many parts of the world as an orna- and the golden lotus logo of Vietnam’s Airline.
mental and as cut flower used as centrepieces for
parties or as religious offerings in India and
Thailand. The dried stalked seed receptacle is Selected References
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Mirabilis expansa
Synonyms
Origin/Distribution
Allionia expansa (Ruiz & Pav.) Kuntze,
Calyxhymenia expansa Ruiz & Pav., Oxybaphus Mauka is indigenous to the Andes in South
expansa (Ruiz & Pav.) Vahl America, distributed naturally in the area of La Paz
(Bolivia), to the North of Quito (Ecuador) and in
Cajamarca (Peru); however, it was also found in
Family Venezuela and Chile (Popenoe et al. 1989; Bermejo
and León 1994; Flores 2003). Its wild ancestors are
Nyctaginaceae found in Peru, Bolivia, Ecuador and Colombia.
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plants. Kampong Publications, Vista. 677 pp The Plant List (2013) Version 1.1. Published on the
Flores H (2003) Andean root and tuber crops: under- Internet; http://www.theplantlist.org
ground rainbows. Hort Sci 38(2):161–167 Vepachedu R, Bais HP, Vivanco JM (2003) Molecular
Klásková T, Fernandéz CE (2011) Mauka – a promise for characterization and post-transcriptional regulation of
fighting with the unbalanced nutrition of high Andean ME1, a type-I ribosome-inactivating protein from
regions. Agric Trop Subtrop 44(1):37–40 Mirabilis expansa. Planta 217(3):498–506
Park SW, Vepachedu R, Owens RA, Vivanco JM (2004) Vepachedu R, Park SW, Sharma N, Vivanvo JM (2005)
The N-glycosidase activity of the ribosome- Bacterial expression and enzymatic activity analysis
inactivating protein ME1 targets single-stranded of ME1, a ribosome-inactivating protein from
regions of nucleic acids independent of sequence or Mirabilis expansa. Protein Expr Purif 40:142–151
structural motifs. J Biol Chem 279(33): Vivanco JM, Savary BJ, Flores HE (1999) Characterization
34165–34174 of two novel type I ribosome-inactivating proteins
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ND, Dafforn M (1989) Lost crops of the Incas. Little expansa. Plant Physiol 119(4):1447–1456
Nymphaea odorata
Nymphaeaceae Agroecology
can be established from seeds in up to 90 cm (8) (Zhang et al. 2003). Compounds 3, 4 and 7
water and the seedlings grow as submerged showed marginal inhibitory effect against fatty
aquatics until they switch abruptly to the produc- acid synthase with IC50 values of 45, 50 and
tion of floating leaves (Richards and Cao 2012). 25 μg/mL, respectively.
Five plant growth inhibitory allelochemicals
from N. odorata were identified as gallic acid,
Edible Plant Parts and Uses myricitrin, myricetin, 1,2,3,4,6-pentagalloyl-D-
glucose and 2,3,4,6-tratragalloyl-D-glucose
Young leaves are used in soups and stews and (Elakovich et al. 1999).
tubers are also edible (Fernald et al. 1958; Tanaka The crude leaf extract of N. odorata exhibited
1976; Kunkel 1984; Facciola 1990). Flower buds weak larvicidal activity against the larvae of the
pickled or eaten as cooked vegetable. malaria vector, Anopheles gambiae, of 10 and
20 % (at 5 % w/v) and 20 and 30 % (at 10 % w/v)
both at 12 and 24 h incubation, respectively
Botany (Oladimeji et al. 2008). The crude leaf extract
and fractions were inactive against the bacterial
An aquatic perennial herbaceous plant with float- and fungal isolates tested.
ing leaves and branched creeping, fleshy rhi- Studies by Deoda et al. (2012) reported that
zomes. The rhizomes are attached by cluster of extracts of the leaf and bulbs exhibited better
adventitious roots arising below the leaf bases; antilithiatic activity than the stem extract of the
they bear circular leaf scars on the upper surface; Dolichos lablab bean extracts.
the rhizomes are brown externally, greyish-white
internally, spongy and are densely covered with
short black hairs. Leaves are circular (10–35 cm Traditional Medicinal Uses
across), cleft at the base, with entire, undulating
margin, shining green and waxy above, reddish The root has been reported to be alterative,
or purplish and more or less hairy on under sur- anodyne, antiseptic, astringent and demulcent
face, petiole slender and cylindrical attached to (Grieve 1971; Lust 1974; Mill 1985; Bown
centre of lower leaf surface (Plate 1). Flowers 1995). A tea made from the roots is used in the
solitary up to 25 cm across, on a grooved pedun- treatment of tuberculosis, chronic bronchial
cle, floating on water surface or raised, with 4 complaints, diarrhoea, dysentery, gastrointestinal
green sepals and 15–40 white or pinkish, lanceo- inflammation, gonorrhoea, vaginal discharge,
late petals with acute apices, 55–120 yellow sta- inflamed glands, mouth sores and to stop bleed-
mens (Plate 1). Fruit an ovoid berry-like capsule, ing (Bown 1995; Foster and Duke 1998). A poul-
1–2 cm across and containing many seeds tice made from the roots is used in the treatment
(2–3 mm long). of swellings, boils, tumours, inflamed skin and
vaginitis (Bown 1995; Foster and Duke 1998).
Moerman (1998) recorded the following
Nutritive/Medicinal Properties ethnomedicinal uses of the plant by American
natives: The Chippewa use pulverized roots for
The following compounds were identified from mouth sores; the Mimac use leaves as cold rem-
fractionation of the ethanol extract of N. odorata: edy, the juice of root is taken for coughs and
nymphaeoside A (1), icariside E(4) (2), kaempferol poultice of boiled roots applied to swellings. The
3-O-α-L-rhamnopyranoside (afzelin, 3), querce- Ojibwe use the root as cough medicine and for
tin 3-O-α-L-rhamnopyranoside (4), myricetin tuberculosis. The Okanagan-Colville place stems
3-O-α-L-rhamnopyranoside (myricitrin, 5), quer- directly onto tooth for toothache. The Penobscot
cetin 3-O-(6″-O-acetyl)-β-D-galactopyranoside apply leaves as poultice on limb swellings. The
(6), myricetin 3-O-β-D-galactopyranoside (7) and Potawatomi used poultice of pounded roots for
myricetin 3-O-(6″-O-acetyl)-β-D-galactopyranoside unspecified ailments.
116 Nymphaeaceae
Nymphaea odorata is an old herbal recipe and stem of Nymphaea odorata and Dolichos lablab
beans. Res J Pharm Biol Chem Sci 3(1):814–819
used in the treatment and/or management of
Elakovich S, Spence S, Yang J (1999) Chapter 4:
ocular, skin, gastrointestinal and urino-genital Phytochemical inhibitors from the Nymphaeceae:
ailments amongst many others (Oladimeji et al. Nymphaea odorata and Nuphar lutea. In: Cutler HG,
2008). Nymphaea odorata has been traditionally Cutler SJ (eds) Biologically active natural products.
CRC Press, Boca Raton, pp 49–56
deemed to be employed as antiseptic, astringent,
Facciola S (1990) Cornucopia. A source book of edible
demulcent, kidney stone, deobstruent, discutient plants. Kampong Publications, Vista. 677 pp
and vulnerary (Deoda et al. 2012). Roots are used Fernald ML, Kinsey AX, Rollins RC (1958) Wild plants
for coughs, mouth sores and for tuberculosis. of Eastern North America, 2nd edn. Harper & Row,
New York
Foster S, Duke JA (1998) A field guide to medicinal plants
in Eastern and Central N. America. Houghton Mifflin
Other Uses Co, Boston. 366 pp
Grieve M (1971) A modern herbal. Penguin. 2 vols. Dover
publications, New York, 919 pp
N. odorata is a common and popular ornamental
Kartesz JT (1999) A synonymized checklist and atlas with
for ponds. biological attributes for the vascular flora of the United
States, Canada, and Greenland. First edition. In:
Kartesz JT, Meacham CA (eds) Synthesis of the North
American Flora, Version 1.0. North Carolina Botanical
Comments Garden, Chapel Hill
Kunkel G (1984) Plants for human consumption. An
A declared aquatic or terrestrial noxious weed annotated checklist of the edible phanerogams and
and/or noxious-weed seed in some states in the ferns. Koeltz Scientific Books, Koenigstein
Lust JB (1974) The herb book. Bantam Books, New York.
USA. Dense infestations may also accelerate the
174 pp
natural siltation process in the shallow bodies of Mills SY (1985) The dictionary of modern herbalism.
water. Thorsons, Wellingborough
Moerman DE (1998) Native American ethnobotany.
Timber Press, Portland. 927 pp
Oladimeji HO, Ubulom PM, Akpabio EI, Etim IE, Nyong
Selected References E (2008) Larvicidal and anti-microbial potentials of
Nymphaea odorata. J Pharmacol Toxicol 3(5):
Bown D (1995) Encyclopaedia of herbs and their uses. 357–362
Dorling Kindersley, London. 424 pp Richards JH, Cao C (2012) Germination and early growth
Deoda RS, Pandya H, Patel M, Yadav KN, Kadam PV, of Nymphaea odorata at different water depths. Aquat
Patil MJ (2012) Antilithiatic activity of leaves, bulb Bot 98(1):12–19
Nymphaea odorata 117
Tanaka T (1976) Tanaka’s cyclopaedia of edible plants of Wiersema JH (1997) Nymphaea. In: Flora of North
the world. Keigaku Publishing, Tokyo. 924 pp America Editorial Committee (eds) 1993+. Flora of
The Plant List (2013) Version 1.1. Published on the Internet; North America North of Mexico. 19+ vols. New York
http://www.theplantlist.org and Oxford
USDA, ARS, National Genetic Resources Program (GRIN) Zhang ZZ, ElSohly HN, Li XC, Khan SI, Broedel SE Jr,
(2014) Germplasm Resources Information Network – Raulli RE, Cihlar RL, Burandt C, Walker LA (2003)
(GRIN) [Online Database]. National Germplasm Phenolic compounds from Nymphaea odorata. J Nat
Resources Laboratory, Beltsville, Maryland. http:// Prod 66:548–550
www.ars-grin.gov/cgi-bin/npgs/html/index.pl
Nymphaea x rubra
Nymphaeaceae
Origin/Distribution
The flowers of the plant are used for ornamental Nutritive/Medicinal Properties
purposes, while rhizomes, young leaves and
peduncle are used as food and vegetable (Devi The proximate nutrient content of various parts
et al. 2015). The species is gaining an importance of N. rubra plant had been determined and found
in the local markets of Northeast India as food to be quite high (Devi et al. 2015). The maximum
material. amount of protein was recorded as 23.88 % of
Rhizome is used as vegetable (Pathak and total dry weight. Proximate nutrient composition
Sarma 2013), the tuber is eaten as a vegetable (per 100 g) of edible tuber was reported as: mois-
(Mohan and Kalidass 2010). The young pedun- ture 81.33 %, crude protein 8.31 g, crude lipid
cles are eaten as vegetable. In India, the fried 5.05 g, crude fibre 3.74 g, ash 3.80 g, N-free
seeds are eaten as puffed grains. The petioles, extractives 79.09 %, energy 1650 kJ/100 g, Na
fruit and roots are cooked as vegetables; seeds are 34.1 mg, K 734 mg, Ca 354 mg, Mg 104 mg, P
eaten raw or roasted; fruit and rootstock are eaten 76.3 mg, Zn 1.64mg, Mn 1.34 mg, Fe 28.14 mg,
raw (Pegu et al. 2013; Patiri and Borah 2007). Cu 1.12 mg, starch 13.52 g, niacin 5.88 mg and
120 Nymphaeaceae
Studies showed N. rubra polysaccharides (NR- The plant is propagated vegetatively by division
PS) to have immunomodulatory effect affecting of rhizome.
the maturation and function of rat dendritic cells
derived from rat bone marrow haematopoietic
cells (Cheng et al. 2012). NR-PS exhibited stimu- Selected References
latory effects on rat dendritic cells and promoted
the secretion of TH1 cytokines. Behera SK, Mohapatra TK, Dash V (2010) Anthelmintic
activity of rhizomes of Nymphaea rubra Linn. Anc Sci
Life 29(3):33–36
Cheng JH, Lee SY, Lien YY, Lee MS, Sheu SC (2012)
Anthelmintic Activity Immunomodulating activity of Nymphaea rubra
Roxb. extracts: activation of rat dendritic cells and
The petroleum ether, methanol and chloroform improvement of the TH1 immune response. Int J Mol
Sci 13(9):10722–10735
extract of N. rubra rhizome exhibited dose- Conrad HS (1905) The waterlilies: a monograph of the
dependent anthelmintic activity against test genus Nymphaea. Carnegie Institution of Washington,
earthworm, Pheretima posthuma (Behera et al. Publication, Washington. 4. 279 pp
2010). The methanolic extract showed remark- Daffodil ED, Mohan VR (2014) In vitro antioxidant activ-
ity of Nymphaea rubra L. rhizome. World J Pharm Res
able anthelmintic activity; at a dose of 5 mg/ml, it 3(4):2178–2189
took 28 min for paralysis and 62 min for worm Devi SA, Thongam B, Handique PJ (2015) Nymphaea
death; at 10 mg/ml it took 17 min for paralysis rubra Roxb. ex Andrews cultivated as an ornamental,
and 51 min for worm death; at 20 mg/ml it took food and vegetable in the North Eastern region of
India. Genet Resour Crop Evol 62:315–320
8 min for paralysis and 27 min for worm death. Dkhar J, Kumaria S, Rao SR, Tandon P (2013) New
The standard albendazole at 10 mg/ml took insights into character evolution, hybridization and
38 min for paralysis and 64 min for worm death. diversity of Indian Nymphaea (Nymphaeaceae): evi-
dence from molecular and morphological data. Syst
Biodivers 11(1):77–86
Gautam S, Rahuja N, Ishrat N, Asthana RK, Mishra DK,
Traditional Medicinal Uses Maurya R, Jain SK, Srivastava AK (2014) Nymphaea
rubra ameliorates TNF-α-induced insulin resistance
The flower is astringent and cardiac tonic and is via suppression of c-jun NH2-terminal kinase and
nuclear factor-κB in the rat skeletal muscle cells. Appl
employed in palpitations of the heart, while the rhi- Biochem Biotechnol 174(7):2446–2457
zomes are used for dysentery and dyspepsia (Khare Gupta PP (1980) Cytogenetics of aquatic ornamentals
2007). Roots are administered for vomiting by rural VI. Evolutionary trends and relationships in the genus
people of Nagaon district, Assam (Sarma and Saikia Nymphaea. Cytologia 45:307–314
Guruge S, Yakandawala D, Yakandawala K (2014)
2010). The Tai-Khamyang of Assam pulverised the Nymphaea rubra Roxb. ex Andrews in Sri Lankan fresh
dried roots to powder and applied the powder in dis- waters. In: Proceedings of international forestry and
eased area for 6–7 days for piles (Sonowal and environment symposium, Sri Lanka. Published by
Barua 2011). In Manipur, north-eastern India, Department of Forestry and Environmental Science,
University of Sri Jayewardenepura. vol 18: P127 (abstract)
crushed rhizome powder is mixed with honey and Juffe Bignoli D (2013) Nymphaea rubra. The IUCN red
used as therapy for bleeding nose, piles and dysen- list of threatened species. Version 2014.2. www.iuc-
tery and as cardiotonic (Yumnam et al. 2012). nredlist.org
Khare CP (2007) Indian medicinal plants: an illustrated
dictionary. Springer, Heidelberg. 900 pp
La-ongsri W, Trisonthi C, Balslev H (2009) A synopsis of
Other Uses Thai Nymphaeaceae. Nordic J Bot 27(2):97–114
Mitra RL, Subramanyam K (1982) Is Nymphaea rubra
Red water lily is used as an ornamental plant in Roxb. ex. Andrews an apomict? Bull Bot Surv India
24:83–86
aquatic landscaping and also in the aquarium Mohan VR, Kalidass C (2010) Nutritional and antinutri-
trade. The cut flowers are used for religious pur- tional evaluation of some unconventional wild edible
poses in India. plants. Trop Subtrop Agrosyst 12:495–506
122 Nymphaeaceae
Pathak U, Sarma DC (2013) A critical study of the biore- Sarma SK, Saikia M (2010) Utilization of wetland
sources of Khaamronga Beel – a wetland for sustain- resources by the rural people of Nagaon district
able development for the people of Thakurkuchi Assam. Indian J Trad Kwon 9(1):145–151
village of Chandrapur area of Assam. Indian J Fund Sonowal R, Barua I (2011) Ethnomedical practices among
Appl Life Sci 3(2):245–251 the Tai-Khamyangs of Assam, India. Ethno Med
Patiri B, Borah A (2007) Wild edible plants of Assam. 5(1):41–50
Geetakhi Printers and Publishers, Guwahati USDA-ARS (2015) National genetic resources program.
Pegu R, Gogoi J, Tamuli AK, Teron R (2013) Germplasm Resources Information Network – (GRIN)
Ethnobotanical study of wild edible plants in Poba [Online Database]. National Germplasm Resources
reserved forest, Assam, India: multiple functions and Laboratory, Beltsville, Maryland. https://npgsweb.ars-
implications for conservation. Res J Agric For Sci grin.gov/gringlobal/taxonomydetail.aspx?415655 .
1(3):1–10 Accessed 7 Mar 2015
Rahuja N, Mishra A, Gautam S, Tamrakar AK, Maurya R, Venu P, Rajakumar TS, Danile P (2003) On the identity of
Jain SK, Srivastava AK (2013) Antidiabetic activity in and sterility in Nymphaea rubra Roxb. ex Andrews
flowers of Nymphaea rubra. Int J Pharm Sci Rev Res (Nymphaeaceae). Rheedia 13:29–34
22(24):121–133 Yumnam RS, Dev CO, Abujam SKS, Chetia D (2012)
Study on the ethnomedicinal system of Manipur. Int
J Pharm Biol Arch 3(3):587–591
Victoria amazonica
Nymphaeaceae
Origin/Distribution
Brazil: Vitória-Regia, Aguapé-Assú, Cará- Victoria amazonica is tropical aquatic. The plant
D’água, Forno-D’água, Forno-De-Jaçanã, roots are best anchored in organically rich loams
Jaçanã, Milho-D’água, Mururê, Nanpé, at the bottom of the bayous. Plants will remain
Rainha-Dos-Lagos, Rainha-Dos-Nenúfare, perennial as long as water temperatures remain at
© Springer International Publishing Switzerland 2016 123
T.K. Lim, Edible Medicinal and Non-Medicinal Plants: Volume 11, Modified Stems, Roots, Bulbs,
DOI 10.1007/978-3-319-26062-4_9
124 Nymphaeaceae
least 24 °C and thrive in calm shallow waters tate, floating, entire and full green adaxially with
(<2 m), full sun, slightly alkaline to neutral the edge turn upwards forming a rim (5–10 cm
pH. Pollination by scarab beetles happens at high), and the basal surface is a coppery colour
night. with a prominent network of flattened ribs radiat-
ing from the petiole and cross ribs, all bearing
sharp prickles (Plates 1–3) (Anonymous 1850;
Edible Plant Parts and Uses Allen 1854). Kunii et al. (2006) reported that the
maximum value in leaf diameter was 2.35 m
In the Brazilian Amazon, Victoria amazonica (4.34 m2 in area) with a mean value of 1.6–1.7 m.
seeds and rhizomes are important items con- Each plant has 10–15 floating leaves and the total
sumed by riverine inhabitants (Piedade and Junk leaf area per plant differed considerably (mini-
2000). The root, stem and seed of the Victoria lily mum value 12.1–17.2 m2, maximum value 27.6–
are all edible, and the species is also used in tra- 31.2 m2). Flower solitary on an aculeate, terete
ditional medicine (Castner et al. 1998). pedicel arising from the rhizome, longer than leaf
The root, stem and seeds are edible. Dried petiole, pear-shaped in bud (Plates 2–3). The
seeds taste like popcorn. Indigenous Brazilians flower is epigynous, fragrant, 30–50 cm across,
make flour from the seeds (Les 2003). hemicyclic, dichlamydeous, bisexual with ves-
pertine anthesis (Rosa-Osman et al. 2011) and
pollinated by scarab beetles. The calyx, consist-
Botany ing of four large and four small spiny sepals
(Kunii et al. 2006), is coriaceous, fleshy and
Victoria amazonica is a rhizomatous aquatic, green becoming reddish purple. Petals are numer-
strongly aculeate, herbaceous perennial (Plate 1) ous, inserted on the throat and torus of the calyx;
with short, thick, tuberous rhizome, which is outer ones are completely reflexed. The petals
brown externally and white inside and turning remained expanded, erect and white all night and
purple when cut; rhizome bears numerous cylin- the next morning; by the evening of the second
drical adventitious roots abounding in air cavi- day, the petals had turned variegated white pink
ties. Leaves are borne on a submerged aculeate, or red. Stamens are numerous, inserted with the
terete petiole 6 m long radiating from the sub- petals on the torus in about three rows; the sta-
merged rhizome; they are huge, orbicular, pel- mens, including the outer staminodia (infertile
Nutritive/Medicinal Properties
Plate 3 Young leaf with coppery basal surface and
flower bud enclosed by four reddish-purple, spiny sepals Two anthocyanins were isolated from Victoria
amazonica leaves and identified as delphinidin
3-O-(2″-O-galloyl-β-d-galactopyranoside) and
stamens) petaloid, middle fertile stamens and cyanidin 3-O-(2″-O-galloyl-β-d-galactopyranoside)
inner staminodia (termed paracarpels by earlier (Strack et al. 1992). Four esters, together with
authors), were erect and formed an approx. free benzyl alcohol, were found as the main
12-mm tunnel into the floral chamber (Seymour components of the fragrance of Victoria (Kite
and Matthews 2006). The filaments are subulate, et al. 1991). The compounds were methyl
erect, outer filaments petaloid, connective long- 2-methylbutyrate, methyl tiglate, benzyl 2-meth-
appendiculate, filaments of the sterile stamens of ylbutyrate and benzyl tiglate. A number of other
the innermost whorl with fleshy appendages; methyl and benzyl esters were found but at much
anthers introrse. Carpels are numerous, crescent lower concentrations. Fifteen compounds were
or horseshoe shaped up to the time of anthesis, isolated from the methanol leaf extract including
and then postgenital fusion occurs forming the stigmasterol, vanillic acids and aristophyll
ovary. The ovary is globose below and concave (Chang et al. 2014).
126 Nymphaeaceae
Na, K, Rb, Cs, AG, Zn, Y, Nb, Se, F, Br, Cl, I Comments
and Ni increased with age; Li, Cu, Be, Mg, Sr,
Cd, Hg, Ce, Si, Ti, Zr, V, Cr, Fe and Al decreased The plant is depicted in the Guyanese coat of
with age; and Ba, B, Ga, Sc, La, P, Bi and S arms.
exhibited no consistent relationship to various Protogyny has been studied in Victoria ama-
growth stages (Cowgill and Prance 1982). zonica (Prance 1974, 1980). The flowers will
However, Ca, Sn, Pb, As, Mo, Mn and Co were open following a dark period of 30 min, begun
low in concentration in young plants, increased not earlier than 4 p.m. (Gessner 1960). The plants
in the pre-flowering stage and then declined with are often pollinated by scarab beetles (Prance and
the stage of the plant. V. amazonica accumulated Arius 1975; Prance 1980).
Na in excess of that encountered in its terrestrial
relatives. It contained rare earths lanthanum (La)
and cerium (Ce) in normal amounts. Rare earths Selected References
of higher atomic numbers were not detectable.
Mg, Ca, Al, Si, S, Fe, Mn, Li and Ce were signifi- Allen JF (1854) Victoria regia or the great water liliy of
America. http://www.victoria-adventure.org/victoria_
cantly more concentrated in the damaged leaf
images/allen_sharp/victoria_regia1.html
sections than in undamaged leaves of the mature Anonymous (1850) Victoria regia. The Gardeners’
plants with unopened buds (Cowgill and Prance Chronicle, pp. unknown
1989). In the case of pre-flowering plants, Na, Castner JL, Timme SL, Duke JA (1998) A field guide to
medicinal and useful plants of the Upper Amazon.
Mg, Ca, Al, Si, S, Cl, Fe and Mn were more prev-
Feline Press, Gainesville. 154 pp
alent in the damaged tissue than in the unattacked Chang MY, Wu HM, Li HT, Li WJ, Chen SJ, Chen CY
leaves of the same plants. In the former case, Na, (2014) Secondary metabolites from the leaves of
K, Cl and P are rapidly lost when damage occurs, Victoria amazonica. Chem Nat Compd
50(5):955–956
while plants that have had a chance to recover
Cowgill UM, Prance GT (1982) Changes in the chemical
from the damage, as in the case of the pre- composition during growth stages of Victoria amazo-
flowering plant leaves, begin to make up the dif- nica (Poeppig) J. de C. Sowerby (V. regia auctt.),
ference in elemental concentration between the Nymphaeaceae. Int Rev Gesamten Hydrobiol
67:235–240
damaged and undamaged leaves. Mineralogical
Cowgill UM, Prance GT (1989) A Comparison of the
investigation showed that the amount of calcite chemical composition of injured leaves in contrast to
(CaCO3), the calcium oxalates weddellite and uninjured leaves of Victoria amazonica
whewellite and the siliceous minerals – low-form (Nymphaeaceae). Ann Bot 64(6):697–706
Duke JA, Vásquez R, Vásquez VM (1994) Amazonian
cristobalite opal, low-form tridymite opal and
ethnobotanical dictionary. CRC Press, Boca Raton,
quartz – was higher in the recently injured leaves 215 pp
than in those that had been injured some time pre- Gessner F (1960) Die Blütenöffnung der Victoria regia in
viously or than in the uninjured leaves. ihrer Beziehung zum Licht. Die Blütenöffnung der
Victoria regia in ihrer Beziehung zum Licht. The
The plant extract has folkloric medicinal use
opening of flowers of Victoria regia in relation to light.
for rheumatism, inflammation and haemorrhoids Planta 54:453–463 (In German)
when mixed with andiroba oil (Duke et al. 1994). Huxley AJ, Griffiths M, Levy M (eds) (1992) The new
RHS dictionary of gardening, (4 vols). MacMillan,
London
Kite G, Reynolds T, Prance GT (1991) Potential
Other Uses pollinator-attracting chemicals from Victoria
(Nymphaeaceae). Biochem Syst Ecol 19(7):535–539
V. amazonica is used as an aquatic ornamental in Kunii H, Sunanisari S, Fukuhara H, Nakajima T, Widjaja
F (2006) Leaf expansion rate and life span of floating
botanical gardens and parks.
leaves in Victoria amazonica (Poepp.) Sowerby grow-
ing in Kebun Raya, Bogor, Indonesia. Tropics
15(4):429–433
Victoria amazonica 127
Les DH (2003) Nymphaeales. In: eLS. John Wiley & Rosa-Osman SMD, Rodrigues R, Mendonça MSD,
Sons Ltd, Chichester. http://www.els.net. doi:10.1038/ Souza LAD, Piedade MTF (2011) Morfologia da
npg.els.0003697 flor, fruto e plântula de Victoria amazonica (Poepp.)
Piedade MTF, Junk WJ (2000) Sustainable management of J.C. Sowerby (Nymphaeaceae). Acta Amazonica
species diversity and primary production of herbaceous 41(1):21–28. [Morphology of flower, fruit and seed-
plants of the Central Amazon floodplain. German- ling of Victoria amazonica (Poepp.) J.C. Sowerby
Brazilian Workshop on neotropical ecosystems – (Nymphaeaceae)]
Achievements and Prospects of Cooperative Research Schneider EL (1976) The floral anatomy of Victoria
Hamburg. Session 6: concepts and paradigms for man- Schomb. J Linn Soc Bot 72:115–148
agement of ecosystem resources. Hamburg Seymour RS, Matthews PGD (2006) The role of ther-
Prance GT (1974) Victoria amazonica ou Victoria regia? mogenesis in the pollination biology of the amazon
Acta Amazon 4(3):6 waterlily Victoria amazonica. Ann Bot 98:
Prance GT (1980) A note on the pollination of Nymphaea 1129–1135
amazonum Mart. & Zucc. (Nymphaeaceae). Brittonia Strack D, Wary V, Metzger JW, Grosse W (1992) Two
32(4):505–507 anthocyanins acylated with gallic acid from the leaves
Prance GT, Arius JR (1975) A study of the floral biology of Victoria amazonica. Phytochemistry 31(3):
of Victoria amazonica (Poepp.) Sowerby 989–991
(Nymphaeaceae). Acta Amazon 5(2):109–139
Diuris semilunulata
None recorded
Edible Plant Parts and Uses
Common/English Names
Botany
Donkey Orchid, Late Leopard Orchid, Spotted
Double-Tail Orchid Diuris semilunulata is a terrestrial, perennial,
geophytic herb. Roots are filamentose with small,
naked, ovoid or cylindrical, often paired, tubers.
Vernacular Names It has 15–25 cm long and 3–4 mm wide, linear,
basal, grass-like, alternate or whorled leaves, that
None recorded is, conduplicate, with 2 leaves in a tuft (Plate 1).
Scape is 20–35 cm high with 3–5-flowered, loose
raceme. Flowers are hermaphroditic, orange,
Origin/Distribution heavily blotched and suffused with brown and
purple. Dorsal sepal is ovate, 6–10 mm long,
This orchid is native to Australia. 7–10 mm wide and erect. Lateral sepals are linear
Nutritive/Medicinal Properties
Other Uses
Synonyms Agroecology
Diuris latifolia Rupp., Diuris oculata F. Muell. This orchid is commonly found in well-drained
ex Lindl., Diuris sulphurea f. immaculata sul- moist soil in semi-shaded sites on slopes of the
phurea Gand., Diuris sulphurea f. tasmanica foothills in open sclerophyll forest and health in
Gand. South Australia, Tasmania, Victoria, New South
Wales and Queensland.
Family
Edible Plant Parts and Uses
Orchidaceae
Starchy root tubers are edible but bland and gluti-
nous and eaten by the aborigines (Anonymous 2010,
Common/English Names 2011; Cribb and Cribb 1987; Low 1989, 1991).
Nutritive/Medicinal Properties
No information on its nutritive value and medici- Plate 1 Long scape with a few flowered raceme
nal properties had been published.
Other Uses
Comments
Selected References
Anonymous (2010) Diuris sulphurea. http://www.yarra-
ranges.vic.gov.au/Residents/Yarra_Ranges_Plant_
Directory/Lower_Storey/Orchids/Diuris_sulphurea
Anonymous (2011) Diuris sulphurea (Tiger orchid).
h t t p : / / bu s h c r a f t o z . c o m / f o r u m s / s h ow t h r e a d .
php?1502-Diuris-sulphurea-%28Tiger-Orchid%29
Cribb AB, Cribb JW (1987) Wild food in Australia, 2nd
edn. Fontana Collins, Sydney, 240 pp
Jones DL (1970) Some thoughts on the taxonomy of the
genus Diuris. Orchadian 3:119–120
Jones DL (1993) Orchidaceae. In: Harden GJ (ed) Flora of
New South Wales, vol 3. New South Wales University
Press, Kensington. 775 pp
Plate 3 Bight yellow flowers with prominent dark- Low T (1989) Bush Tucker – Australia’s wild food har-
coloured markings vest. Angus & Robertson, North Ryde, 233pp.
Low T (1991) Wild food plants of Australia. Angus &
Robertson, North Ryde. 240 pp
Pridgeon AM, Cribb PJ, Chase MW, Rasmussen FN (eds)
(2001) Genera Orchidacearum: Volume 2 Orchidoi-
deae (Part one). Oxford University Press, Oxford/
New York, 438 pp
Pterostylis curta
Common/English Names
Botany
Maroonhood
Pterostylis pedunculata is a terrestrial perennial
herb, 25 cm high, arising from small round
Vernacular Names tubers to form large colonies. It has a ground-
hugging rosette of 2–6, shortly petiolate, thick,
None recorded dark green, ovate to oblong, 10–65 mm long by
6–18 mm wide leaves, with entire or wrinkled
margins (Plates 1 and 2). Scape is 20–25 cm
Origin/Distribution with a single erect flower, that is, 15 mm long,
white and green with reddish-brown to black
The species is indigenous to Australia. suffusions in the galea. Central sepal and petals
are united into a galea (hood) over the column wide, dark reddish brown, straight and apex
and labellum (lip) (Plates 2, 3 and 4). Apex of obtuse.
galea is horizontal. Dorsal sepal is acute to sub-
acute. Lateral sepals are erect, loosely embrac-
ing the hood, leaving a broad V-shaped lateral Nutritive/Medicinal Properties
gap; free points are filiform, 15–30 mm long,
divergent and recurved. Petals are narrow and No information on its nutritive value and medici-
acute. Labellum is ovate, 4–7 mm long by 3 mm nal properties had been published.
138 Orchidaceae
Other Uses
Comments
Selected References
Anonymous (2011) Pterostylis pedunculata. http://www.
yarraranges.vic.gov.au/Residents/Yarra_Ranges_
Plant_Directory/Lower_Storey/Orchids/
Pterostylis_pedunculata
Jones DL (1993) Orchidaceae. In: Harden GJ (ed) Flora of
New South Wales, vol 3. New South Wales University
Press, Kensington. 775 pp
Steenbeeke G (2001) List of plant species from northern
NSW that may be used as food plants. http://brg.cma.
nsw.gov.au/uploads/MurriFoodPlants.pdf
wild forms is found in the central region of Peru thrives best in light, well-drained and rich soil in
(latitude 10°S) and northern Bolivia (latitude full sun. Oca requires day length shorter than
20°S). Oca is grown in greatest abundance in the 12 h to stimulate tuber formation; longer day
highlands of Ecuador, Peru, Columbia and length periods promote foliage growth.
Bolivia, although it is found as far north as
Venezuela and as far south as Chiloe Island in
Chile (Popenoe et al. 1989). Of all Andean root Edible Plant Parts and Uses
and tuber crops, oca is presently second only to
potato in area planted within the Central Andean Oca tubers are boiled, baked, roasted, fried,
region. Oca was introduced into Europe in the mixed fresh with salads, added to stews and
last century but did not establish as a permanent soups and pickled in vinegar or served as a sweet,
vegetable crop. In recent decades, oca has estab- either plain or candied (Arbizo and Tapia 1994;
lished as a popular commercial vegetable crop King and Gerhoff 1987; Popenoe et al. 1989;
in New Zealand, where it called yam (Popenoe Bradbury and Emshwiller 2011). In the Andes, a
et al. 1989). few genotypes types are eaten raw. In Mexico,
Earlier results of studies with nuclear-encoded, oca is commonly sprinkled with salt, lemon and
chloroplast-expressed glutamine synthetase sug- hot pepper, and eaten raw. It is also made into
gested that Oxalis picchensis might be a progeni- bottled preserves (often in vinegar). Ocas that are
tor of oca (Emshwiller and Doyle 2002). consumed fresh belong to the category wayk’u
Subsequent AFLP (amplified fragment length (boiling) or misk’i (sweet, delicious). Aymara
polymor phism) data of this species, as well as speakers refer to ocas for processing as luk’i and
different populations of wild, tuber-bearing those for fresh consumption as q’eni. In the
Oxalis found in Lima Department, Peru, were Andes, Oca is first sun-dried for a few days, to
relatively divergent from O. tuberosa (Emshwiller make it sweeter and then parboiled, roasted or
et al. 2009). Results from all analytical methods prepared as pachamanca (meat roasted in a hole
suggested that the unnamed wild, tuber-bearing in the ground). The dried, frozen tuber is called
Oxalis found in Bolivia and O. chicligastensis in khaya. If it is washed after freezing, a whiter
NW Argentina were the best candidates as the product called okhaya is obtained which is con-
genome donors for polyploid O. tuberosa. sidered to be of superior quality. The flour of the
latter is used to make porridges and desserts.
Bitter varieties are almost always converted into
Agroecology dry products (cavi or caya), during which the bit-
terness disappears to leave bland-tasting products
Oca is widely grown as a staple crop in the that can be stored without refrigeration. The bit-
Andean highlands due to its tolerance of mar- terness can also be removed by soaking and
ginal soil, acidities between pH 3.5–7.8, high repeatedly freeze-drying the tubers. Oca tubers
altitude, harsh climate and ease of propagation also have potential for producing superior quality
(Popenoe et al. 1989; Arbizu and Tapia 1994). In starch for the food industry or alcohol.
its native range, it is cultivated from 2800 to Sensory evaluation studies in New Zealand
4000 m altitudes. In New Zealand, it is grown found that for raw oca, panellists preferred bright
near sea-level. Oca flourishes in a climate of low red tubers, but size was also important (Sangketkit
cool temperatures from 4 to 18 °C and mean et al. 2000). Flesh colour, bitterness and meali-
annual precipitation of 570–2000 mm uniformly ness were important variables which described
distributed throughout the growing season. It tol- the overall acceptability of steamed oca, while
erates light frost but its top growth is severely only flesh colour and bitterness were important
damaged by sub-zero temperatures. Temperatures variables for baked oca. Cultivars which had flesh
above about 28 °C cause the plant to wilt and its colour described as ‘bright yellow’, flavour as
leaves to die and tuber production reduced. It ‘no bitter taste’ and a ‘very slightly mealy’ tex-
Oxalis tuberosa 141
ture were more preferred. Cultivars with higher and form tubers. The tubers are claviform-ellip-
yellow/blue (b∗) and chroma (c∗) values for soid and cylindrical, 3–15 cm long by 3 cm wide,
cooked tuber skin and flesh colours were more stubby-looking, with buds on the whole surface,
acceptable. Martin et al. (2005) found that culti- and variegated in colour: white, yellow, red and
vars with red skin, bright yellow flesh colour, a purple. Leaves trifoliate on 2–9 cm long petioles,
slightly mealy texture and a sweet taste were leaflets, heart-shaped, clover-like and green.
preferred. Inflorescence with 4–5 small flowers. Calyx with
five pointed green sepals. Corolla with five
obovate, clawed petals fused at the base and with
Botany radiating purple striations at the basal part.
Stamens ten. Flowers are tristylous: short-styled
Oca is a compact, perennial, tuberous herb, erect flowers with two tiers of stamens above the style;
becoming decumbent or prostrate towards matu- mid-styled flowers with a tier of stamens both
rity, usually 20–30 cm high, with cylindrical, above and below the styles and long-styled flow-
succulent stems that vary in colour from yellow ers with two tiers of stamens below the styles.
and green to a purplish red (Plates 1–2). Under
long days, the stolons grow as above-ground
stems; under short days, they penetrate the soil Nutritive/Medicinal Properties
Oca tuber was reported to have a nutritional Hernández-Lauzardo et al. (2004) reported
value equivalent to or better than potato; on aver- that oca starch possessed an apparent amylose
age, oca contained 84.1 % moisture, 1.1 % protein content of 33 %, similar to maize starch used as
and 0.6 % lipid (Popenoe et al. 1989). Fourteen control, with a granule size between 25 and
percent of its nutrient content comprised carbohy- 50 μm with oval and elliptical shapes and A-type
drates with sucrose as the major sugar (Gross X-ray diffraction pattern. The gelatinization tem-
et al. 1989). Chemical composition of oca tubers perature of oxalis starch was 64.0 °C, that was
contains protein 6.84 %, lipid 0.72 %, soluble lower than the one determined in maize starch
fibre 0.29 %, insoluble fibre 6.85 %, total sugar (73.0 °C), with an enthalpy value of 12.2 J/g,
23.28, starch 56.82 %, mineral residues 4.48 % which was similar to that of maize starch. Both
and moisture 86.23 % (Valcárcel-Yamani et al. oxalis and maize starch pastes behaved as weak
2013). Chemical composition of oca contains viscoelastic systems with the elastic modulus
starch 99.21 %, amylose 27.60 %, mineral resi- (G′) predominating over the viscous character
dues 0.15 % and moisture 10.93 %. Oca starch (G″). Heating (gelatinization) caused a more pro-
granules were ellipsoid, oval, conical, pear-shaped nounced enhancement in the structure of the
and prismatic forms: ellipsoids and oval granules maize starch pastes than in that of oxalis starch
with lengths up to 54.30 μm. The physical, chemi- pastes. They concluded that due to its physico-
cal and functional characterization of starches chemical, functional and rheological properties,
from Andean tubers oca, mashua and ulluco sug- oxalis starch could be suitable for testing its use
gested that these starches could be used in food in the cosmetic and food industry.
systems and other industrial applications, in prod- Total phenolic content of oca tubers ranged
ucts that require easy cooking, hot high viscosity, from 0.71 to 1.32 mg/g with the purple coloured
stability under refrigeration and do not need to be varieties having higher content and the yellow
frozen. The absence of protein in the isolated coloured varieties lower phenolic content
starches indicated the utility of these starches for (Campos et al. 2006). Total carotenoid content
preparing syrups with high glucose content. The ranged from 2 to 25 μg/g in oca, moisture content
starches were found to cook easily and to have a 80–85 % and lipophilic antioxidant in oca tubers
high degree of swelling and solubility, high vis- ranged from 69 to 320 μg/g TE. For oca pig-
cosity, low stability to stirring and cooking or mented tubers, the total anthocyanin content
mechanical action and a low tendency towards ranged from 0.14 to 1.3 mg/g; anthocyanin was
retrogradation. These starches showed high clar- concentrated in the tuber skin.
ity, but with high syneresis when subjected to Eight different anthocyanins were found in the
freeze-thaw cycle. Oca starch granules were oval- coloured tubers of isla oca; the major compounds
shaped with granular sizes between 20 and 55 μm, were malvidin 3-O-glucoside and malvidin
amylose content of 18.4 %, amylopectin with 3,5-O-diglucoside; the rest were petunidin
average chain length of 22.4 and β-amylolysis 3,5-O-diglucoside, peonidin 3,5-O-diglucoside,
limit of 64.5 %, B-type X-ray diffraction pattern, petunidin 3-O-glucoside, delphinidin 3-O-glucosides
gelatinisation enthalpy of 14.6 J/g and peak tem- and peonidin3-O-glucoside (Alcalde-Eon et al.
perature of the endothermic DSC (differential 2004).
scanning calorimetry) transition was 55.9 °C Caffeic, vanillic and cinnamic acid deriva-
(Santacruz et al. 2002). Santacruz et al. (2003) tives, flavan-3-ols and flavones derivatives, were
found that oca starch gel stored at 4 °C showed a identified as the main non-anthocyanin phenolic
high increase in G′ during the first day of storage. compounds in the aqueous tuber fraction of two
A decrease in pH from 6.5 to 4.0 produced a loss different coloured oca genotypes (Chirinos et al.
of structure in the starch gel, as was showed by 2009). Acid hydrolysis revealed the presence of
the reduction of G′. Storage at freezing tempera- vanillic, caffeic and cinnamic acids and malvidin
ture (−20 °C) produced higher changes in G′ than in the aqueous fraction. The ethyl acetate tuber
refrigeration conditions. fraction was composed mainly of caffeic and cin-
Oxalis tuberosa 143
namic acid derivatives as well as flavan-3-ols, but markedly decreased contents of starch and
flavones and flavanone derivatives. The flavan-3- organic acid. The chemical composition (range,
ols, flavones and flavanones detected in both per 100 g edible matter) reported for fresh and
fractions corresponded to bound forms of cate- sunned tubers was respectively as follows: dry
chin, luteolin and apigenin and naringenin, matter 10.4–14.3 g, 16.1–20 g; starch 5.2–9.2 g,
respectively. The aqueous tuber fractions were 2.2–6.5 g; soluble solids 6.2–92, 6.7–9.7 oBrix;
the major contributors to the ABTS antioxidant total sugars 2119–3635 mg, 2910–5870 mg;
capacity (77–82 %). sucrose 1208–1826 mg, 1901–3955 mg; fructose
Storage at 4 °C substantially extended the 477–1085 mg, 553–1217 mg; glucose 433–
storage life of oca tubers held for 20 weeks (Kays 1025 mg, 456–988 mg; organic acids 2567–
et al. 1979). Respiratory rate (4 mg CO2/kg/h at 3385 mg, 937–1970 mg; glutarate
4°), fresh weight and colour loss, and sprouting 1071–1578 mg,121–516 mg; malate 636–
were retarded. Composition changes of the tubers 1088 mg,139–660 mg; oxalate 306–539 mg,251–
(% reducing sugars, non-reducing sugars, total 451 mg; succinate 270–375 mg, 294–364 mg and
sugars, starches, total carbohydrates, total nitro- tartarate 57–138 mg, 56–191 mg. Twelve South
gen and non-protein nitrogen) were less pro- American and two New Zealand cultivars of oca
nounced at 4 °C than at 21 °C. Tubers stored at were found to contain total oxalate levels of
21 °C underwent substantial changes in composi- 80–194 mg/100 g wet matter (WM) in raw tubers
tion, the majority of which tended to coincide (Sangketkit et al. 2001). Oxalate levels ranged
with sprout formation. The mean dry matter con- from 77 to 220 mg/100 g WM for boiled and
tent of freshly harvested oca tubers was steamed tubers, while the levels of oxalate found
14.3 g/100 g fresh weight (FW), and after storage in baked tubers were significantly increased
for 6 weeks at 16.4 °C was 14.6 g/100 g FW when compared to the raw tubers. The oxalate
(Savage et al. 2008). The mean soluble oxalate content of the baked tubers ranged from 164 to
content of freshly harvested tubers was 335 mg/100 g WM. Total calcium content of the
162.1 mg/100 g FW, and, after 6 weeks storage, raw tubers ranged from 7.5 to 15.5 mg/100 g
173.5 mg/100 g FW. WM; cooking had a little effect on the calcium
Six organic acids had been reported in oca content. Among cultivars mean oxalate/calcium
tubers: ascorbic, malic, tartaric, succinic, glutaric ratio of the raw, boiled and steamed tubers ranged
and oxalic acids (Hermann and Erazo 2000; Ross from 2.5 to 9.9. Baked tubers had a significantly
et al. 1999; Bradbury and Emshwiller 2011). Of increased oxalate/calcium ratio (mean for all cul-
these oxalic acid is most deleterious to human tivars 9.5). Albihn and Savage (2001a, b) found
health (Ross et al. 1999; Albihn and Savage that oca tuber contained moderately high levels
2001a, b; Sangketkit et al. 2001). King (1988) of oxalate which existed in soluble form only.
reported low levels of 20.3–50.3 mg/100 g FW They found that oca tuber skin contained signifi-
oxalic acid in the tubers; in contrast, Ross et al. cantly higher oxalate level than the flesh 7.3 g/kg
(1999) reported levels of 92–221 mg/100 g FW in fresh weight (FW) compared to 1.7 and 1.4 g/kg
12 South American and two New Zealand culti- FW in the outer and inner flesh, respectively).
vars of oca. Levels of soluble and total oxalate The highest concentration was found in the skin
extracted from the tubers were not significantly of the pinkish-red cultivar (10.9 g/kg FW). All
different, suggesting that no calcium oxalate was cooking methods appeared to cause a migration
formed in the tuber. Hermann and Erazo (2000) of oxalate from the skin to the underlying flesh.
reported the highest oxalate value of 306– On a fresh weight basis, baking significantly
539 mg/100 g FW in freshly harvested tubers and increased the oxalate concentration in the whole
251–451 mg/100 g in sunned tubers. Sunning tuber, whereas boiling decreased the concentra-
reduced oxalate levels by an average of 26 %. In tion and steaming had no significant effect.
comparison to fresh tubers, sunning increased Boiling might therefore be a better way of cook-
dry matter content, soluble solids and total sugars ing oca than baking when a low intake of oxalate
144 Oxalidaceae
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Piper methysticum
narcotic substance from dried kava stems and The content (weight %) of six major kavalac-
roots, and called it marindinin which later was tones in the dried kava roots was reported as:
found to be identical with dihydrokawain of kavain 2.58 %, dihydrokavain 1.37 %, methysti-
Borsche. The concentrations of desmethoxyyan- cin 1.82 %, dihydromethysticin 1.89 %, yangonin
gonin, yangonin, dihydromethysticin, methysticin, 1.73 %, desmethoxyyangonin 0.81 % analysed
dihydrokawain and kawain in kava ether extract in Hawaii (Young et al. 1966); kavain 1.90 %,
were determined (Young et al. 1966). Dutta et al. dihydrokavain 2.37 %, methysticin 2.12 %, dihy-
(1972) isolated yangonin and methysticin from dromethysticin 1.12 %, yangonin 1.21 %, desme-
kava roots. Recoveries of known compounds were thoxyyangonin 0.59 % analysed in Fiji (Duve
80–85 % for yangonin and 90–95 % for all others. 1981); kavain 2.30 %, dihydrokavain 3.28 %,
Fresh kava root stock contained 80 % water, while methysticin 2.06 %, dihydromethysticin 1.60 %,
dried rootstock consists of approximately 43 % yangonin 1.16 %, desmethoxyyangonin 0.91 %
starch, 20 % fibre, 12 % water, 3.2 % sugars, 3.6 % analysed in Vanuatu (Lebot and Lévesque 1996).
proteins, 3.2 % minerals and 15 % (3–20 %) kava- The composition of the major kavalatones in kava
lactones (Lebot et al. 1992). Kavalactone content root, stem and leaf were reported respectively by
was reported to be greatest in the roots and to Smith (1983) askavain (34.5, 0.6, 2.5 %); dihy-
decrease higher up the plant. Relative concentra- drokavain (17.1,23.2,69.8 %); methysticin (20.8,
tions of 15 %, 10 % and 5 % have been observed 13.9, 0.8 %); dihydromethysticin (5.3, 59.6,
in the root, stumpand basal stems, respectively. 22.5 %); yangonin (0.8,0.8, 1.2 %) and desme-
Kavalactones (amounting to %) had been reported thoxyyangonin (21.6, 1.8, 3.0 %).
in all parts of the kava plant comprising the major The first compound isolated from Piper
ones: kavain (1.8 %), methysticin (= kavahine, methysticum was methysticin (Cuzent 1981) also
kavakin, kavatin, kanakin) (1.2 %), demethoxy- known as kavatin, kavahin and kanakin. The
yangonin (1 %), yangonin (1 %), dihydrokavain IUPAC has recommended the name 5,6-dihydro-
(0.6 %), dihydromethysticin (0.5 %) and traces of 4-methoxy-6-[3′,4′-(methylenedioxy)styryl]-2H-
11,12-dimethoxyhydrokavain, 11-hydroxy12- pyran-2-one but chemical abstracts indexing has
methoxykavain, 11-methoxy-nor-yangonin, adopted the latter path, using 5-hydroxy-3-
11-methoxy-yangonin and the two ethylketones methoxy-7-[3,4-methylenedioxyphenyl]-2,6-
cinnamoylacetone and methylendioxy-3,4- hepta-dienoic acid- lactone. The ethylenic
cinnamoylidenacetone (Shulgin 1973; Young et al. dihydroanalog dihydromethysticin was isolated
1966). almost 50 years later by Winzheimer (1908).
Piper methysticum 151
Nolting and Kopp (1874) isolated yangonin. The A total of 19 kavalactones namely 10-
most extensive studies on the constituents of kava methoxyyangonin; hydroxykavain; dihydro-
plant were conducted by Borsche and coworkers 5,6-dehydrokavain; 7,8-dihydrokavain; 7,8-
in 13 papers wherein they reported the isolation dihydroyangonin; kavain; 7,8-dihydro-5-
and structure of two compounds kawain (also hydroxykavain; 5,6-dihydroyangonin; 11-
known as kavain and gonosan) and dihydro- h y d r oxy-12-methoxydihydrokavain;
kawain and the structural elucidation of the crys- 11-methoxyyangonin; 5,6-dehydrokavain (des-
talline components methysticin, dihydromethysticin methoxyyangonin); 5,6,7,8-tetrahydroyango-
and yangonin (Borsche 1927; Borsche and nin; methysticin; dihydromethysticin; 11,12
Bodenstein 1929; Borsch and Blount 1930, 1932, dimethoxydihydrokavain; yangonin; 11-methoxy-
1933; Borsche and Gerhard 1914; Borsch and 12-hydroxydehydrokavain; 11-hydroxyyango-
Lewinsohn 1933; Borsche et al. 1927a, b, 1929a, nin; 5,6-dehydromethysticin were identified in
b; Borsche and Peitzsch 1929a, b, 1930; Borsche kava roots (Hänsel 1968; Lebot and Cabalion
and Roth 1921; Borsche and Walter 1927). 1988; He et al. 1997; Whittaker et al. 2008;
Yangonin (= 4-methoxy-6-(p-methoxy-β-styryl)- Dharmaratne et al. 2002; Xuan et al. 2008;
α-pyrone) was not so recognised by Borsche and Teschke 2011; WHO 2007), with flavokavains A,
co-workers (Borsche and Gerhard 1914; Borsche B and C (He et al. 1997); kava lactones (Hänsel
and Bodenstein 1929). They had postulated it as et al. 1968) identified in kava roots included:
a γ-pyrone, although they had correctly inter- kavain; 7,8-dihydrokavain; 5,6-dehydrokavain;
preted the structures of all other substances iso- yangonin; 5,6,7,8-tetrahydroyangonin; methysti-
lated from kava (Borsche et al. 1927a, b; Borsche cin; dihydromethysticin; 5,6-dehydromethysticin;
and Peitzsch 1929a, 1930). Yangonin served as 5,6-dihydroyangonin; 7,8-dihydroyangonin; 10-
the basis for two additional substances the meta- methoxy-yangonin; 11-methoxy-yangonin; 11-
methoxy analogue 11-methoxyyangonin (Hänsel hydroxy-yangonin; hydroxykavain; 11-methoxy-
and Klaproth 1966) and the phenolic counterpart 12-hydroxy-dehydrokavain (Lebot and Cabalion
11-methoxy-nor-yangonin (Hänsel et al. 1966c) 1988). Major compounds identified in Kava
isolated from Kava-kava, both of which had been extracts by low-resolution GC/MS included
established synthetically as possessing the kava lactones like kavain, dihydro-and dehydro-
α-pyrone ring. The completely conjugated ana- kavains, yangonin and tetrahydroyangonin,
logue, 5,6-dehydromethysticin, was identifed as methysticin and dihydromethysticin (Nerurkar
a component of kava-kava by Hänsel (1968). The et al. 2004; Lopez-Avila and Benedicto 1997).
following kavalactones were isolated from kava Other kavalactones were identified as minor
roots: 5,6,7,8-tetrahydro-yangonin (Achenbach components including 1-methoxyyangonin
et al. 1971); 11-hydroxy-12-methoxy-dihydro- (Hänsel and Klaproth 1966), 11-metho
kawain and 11,12-dimethoxy-dihydrokawain xynoryangonin and 5,6-dehydromethysticin
(Achenbach et al. 1972). Two yellow pigment (Hänsel et al. 1966b); 1-methoxyyangonin
materials, flavokawin A and B, were isolated (Hänsel and Klaproth 1966), 11-methoxynoryan-
from kava rhizomes (Hänsel et al. 1961). The gonin and 5,6-dehydromethysticin (Hänsel et al.
structures of these were established by synthesis 1966b); 10-methoxyyangonin, 11-hydroxyyan-
to be substituted chalcones bearing an obvious gonin and 11-methoxy-12-hydroxydehydroka-
biogenetic relationship to the styrylpyrones vain (Duve 1981; He et al. 1997). Additionally,
(Hänsel et al. 1963). Hänsel and Schulz (1973) three hydro-derivatives of yangonin were identi-
synthesized 5,6,-cis-kavain 5-ol and 5,6-trans- fied from kava plant. These include 5,6-dihydroy-
kavain-5-ol. Flavokawin A was elucidated as angonin (Hänsel 1968; Duve 1981),
2′-hydroxy-4-4′-6′-trimethoxychalcone and fla- 7,8-dihydroyangonin (Duve 1981) and 5,6,7,8-tet-
vokawin B as 2′-hydroxy-4′-6′ dimethoxychal- rahydroyangonin (Achenbach et al. 1971; Hänsel
cone. The flavokavins A, B and C were reported 1968). Hydroxylated α-pyrone derivatives, includ-
to have a different skeleton (Duve 1976; Hänsel ing cis-5-hydroxykavain (Duve 1981) and
et al. 1963; Dutta et al. 1973, 1976). 5-hydroxydihydrokavain (Achenbach and
152 Piperaceae
Wittman 1970), had also been isolated from P. and 39 months. After 39 months of storage 93.9 %
methysticum as minor components. of dihydrokavain, 81.6 % of kavain, 72.4 % of
The kavalactone, 11-methoxy-5,6-dihydro- dehydrokavain, 54.9 % of tetrahydroyangonin,
yangoninand eight previously reported analogs 25.8 % of dihydromethysticin, 32.1 % of yangonin
along with four other aromatic compounds were and 29.5 % of methysticin had deteriorated in the
isolated from kava root extracts (Ranjith et al. powdered root samples.
2002). Nine kava lactones were detected from Concentrations of total kava lactones
kava roots using GC-MS including desmethoxyy- (methysticin, dihydromethysticin, kawain, dihy-
agonin, kavain, 7,8-dihydrokavain, hydroxyka- drokawain, yangonin and desmethoxyyango-
vain, yagonin, 5,6,7,8-tetrahydroxyyagonin, nin), ranged between 135 and 0.035 mg per
methysticin, dihydromethysticin and 11-hydroxy- serving in 10 beverage products, two chocolate
12-methoxydihydrokavain (Xuan et al. 2006). products, three unbrewed tea products, a drink
Quantities of desmethoxyyagonin, kavain, mix product tested and between 40 and 61 mg
7,8-dihydrokavain, yagonin, methysticin and per serving for the three dietary supplement
dihydromethysticin detected were 4.3, 6.9, products tested using LC-UV and LC-MS anal-
18.6, 5.7, 1.4 and 5.4 mg/g of dry weight, respec- yses (de Jager et al. 2004). One percent of kava
tively. Eighteen kava lactones, cinnamic acid bor- extract could be detected in food supplements
nyl ester and 5,7-dimethoxy-flavanone, and seven using liquid chromatography-atmospheric pres-
compounds, including 2,5,8-trimethyl-1-naphthol; sure chemical ionisation tandem mass spec-
5-methyl-1-phenylhexen-3-yn-5-ol; 8,11-octa- trometry, corresponding to approximately
decadienoic acid-methyl ester; 5,7-(OH)(2)-4′- 0.05–0.2 mg/g of the individual kava lactones
one-6,8-dimethylflavanone; pinostrobin chalcone; kavain, dihydrokavain, yangonin, desmethoxyy-
and 7-dimethoxyflavanone-5-hydroxy-4′, were angonin, methysticin and dihydromethysticin
identified from kava roots (Xuan et al. 2008). (Bobeldijk et al. 2005). Reliable quantification
Glutathione (26.3 mg/g) was found in the water was obtained from concentrations of
extract. Dihydro-5,6-dehydrokavain (DDK) was 0.25–1 mg/g, depending on the compound.
present at a higher level than methysticin and des- Reliable quantification was obtained from con-
methoxyyagonin, indicating that DDK to be a centrations of 0.25–1 mg/g, depending on the
major constituent of kava roots. compound. Gautz et al. (2006) found the combi-
Of 18 kavalactones or kava pyrones found in nation of near-infrared reflectance spectroscopy
kava, kawain, dihydrokawain, methysticin, and partial least-squares methods to be a conve-
dihydromethysticin, yangonin and desmethoxyy- nient, versatile and rapid analytical tool for
angonin were the six major kavalactones determination of kavalactones in dried kava
(Whittaker et al. 2008). The following kavalactone powder.
compounds were isolated from kava roots: Major and trace constituents of kava resin
kavalactones:10-methoxyyagonin; hydroxykavain; were identified using methane negative ion
dihydro-5,6-dehydrokavain; 7,8-dihydrokavain; chemical ionization (NICI) mass spectrometry
7,8-dihydroyagonin; kavain; 7,8-dihydro-5- (Duffield and Lidgard 1986; Duffield et al. 1986).
hydroxykavain; 5,6-dihydroyagonin; 11-hydroxy- Several of the kava lactones (kawain, methysti-
12-methoxydihydrokavain; 11-methoxyyagonin; cin, desmethoxyyangonin and yangonin) pro-
desmethoxyyagonin, 5,6,7,8-tetrahydroyagonin; duced abundant molecular anions when analysed
methysticin; dihydromethysticin; 11,12-dimethy- by methane NICI mass spectrometry, in contrast
oxydihydrokavain; yagonin; 11-methoxy-12-hy- to their 7,8-dihydro analogues which ionized
droxydehydrokavain; 11-hydroxyyagonin; and with poor efficiency to yield weak [M − H]−
5,6-dehydromethysticin (Xuan et al. 2008). Duve anions. Several new trace compounds formally
and Prasad (1983) investigated the stability of kav- resulting from decarboxylation of the 4-methoxy-
alactones in powdered kava root stored in screw- 2-pyrone ring system in the known compounds
capped glass bottles at room temperature for 22, 36 were also found. A simultaneous HPLC separa-
Piper methysticum 153
tion of the enantiomers of kavain, dihydrokavain, For ground kava (250–500 μm), 2 h of sub-
methysticin and dihydromethysticin as well as critical water extraction was required for a com-
the achiral dienolides yangonin and desme- plete extraction of kava lactones at 100 °C, while
thoxyyangonin was carried out on a ChiraSpher at 175 °C, 20 min were sufficient (Kubátová et al.
NT column (Boonen et al. 1997; Häberlein et al. 2001). For a complete extraction of the unground
1997). Using reverse-phase high-performance (shredded) kava, the time of extraction was
liquid chromatographic method, six major kava- extended to 40 min at 175 °C. Boiling for 2 h and
lactones [methysticin, dihydromethysticin extraction with Soxhlet apparatus for 6 h, both of
(DHM), kavain, dihydrokavain (DHK), deme- which employed water at atmospheric pressure,
thoxyyangonin (DMY) and yangonin] in Piper produced yields 40–60 % lower than those
methysticum (kava) were simultaneously deter- obtained with subcritical water. With unground
mined (Shao et al. 1998). The detection limits kava, 40 min of subcritical water extraction
(S/N = 3) for methysticin, DHM, kavain, DHK, yielded essentially the same recoveries of lac-
DMY and yangonin are approximately 0.5, 1.1, tones as 18 h of sonication with acetone, methy-
0.7, 1.1, 0.6 and 0.6 μg/ml, respectively. The lene chloride or methanol. Kava root extracted in
average recoveries are 100.2 % for methysticin, acetone yielded 100 % kavalactones, 96 % etha-
100.6 % for DHM, 100.0 % for kavain, 100.3 % nol yielded 100 % kavalactones, 25 % ethanol
for DHK, 98.9 for DMY and 98.2 % for yango- yielded 15 % kavalactones and water yielded
nin. A rapid micellar electrokinetic chromatogra- 2.97 % kavalactones (Denham et al. 2002).
phy method with diode-array detection was used Extraction rates also varied depending on the
to separate kavalactones kavain, dihydrokavain, temperature at which the products were prepared.
methysticin, dihydromethysticin, yangonin and Analysis of flavokavins in extracts of 172 sam-
demethoxyyangonin from dry kava extracts ples originating from four cultivars groups
(Lechtenberg Lechtenberg et al. 1999). (noble, medicinal, 2-days and wichmannii) indi-
Three kava lactone constituents of Piper cated that the ratio flavokavin B /kavalactones
methysticum, namely, kawain, methysticin and was much higher in 2 days (0.39) and wichman-
desmethoxyyangonin, were separated and identi- nii (0.32) compared to nobles (0.09) and medici-
fied by reverse-phase HPLC using superheated nal cultivars (0.10) (Lebot et al. 2014). For each
deuterium oxide as the mobile phase and on-line group, the ratios of flavokavins/kavalactones did
1H-NMR detection (Chienthavorn et al. 2005). not change significantly between roots, stumps or
Meissner and Häberlein (2005) used simultane- basal stems and among clones, indicating that
ous HPLC for the separation of six major kavapy- they were genetically controlled.
rones and the flavokavins A–C in an ethanolic
extract of Piper methysticum. Flavokavins A–C
contents of 0.62 mg/100 mg, 0.34 mg/100 mg Phytochemicals – Alkaloids/
and 0.14 mg/100 mg ethanolic kava extract was Flavonoids/Miscellaneous
determined respectively. Compounds identified Compounds
in commercial kava extracts by gas chromatogra-
phy (GC) and quadrupole time-of-flight mass There had been reports of isolation of alkaloids,
spectrometry (QTOFMS) with an electron flavonoids – flavokavins, sterols, organic acids,
ionization source: 7,8 dihyrokavain; kavain; aliphatic alcohols and other compounds from
5,6-dehydrokavain; 5,67,8-tetrahydroyangonin; kava plant. Three flavokavins (also known as fla-
7,8-dihydromethysticin; yangonin; methysti- vokawins or flavokavains), designated A, B and
cin; bornyl cinnamate; dihydroxy- C, had been identified from kava root (Dutta and
methoxyphenyl-phenyl-propen-one; hydroxyl- Ray 1973; Dutta et al. 1973, 1976; Hänsel et al.
dimethoxyphenyl-3-phenyl-2-propen-1-one; and 1961, 1963). Structures of these compounds had
hydroxyl-dimethoxyphenyl-3-methoxyphenyl-2- been confirmed by synthesis (Hänsel et al. 1963;
propen-1-one (Lopez-Avila and Yefchak 2009). Dutta et al. 1976; Dutta and Som 1978). Two
154 Piperaceae
and Wee1, and caused cyclin B1 protein accumu- with minimum toxicity. Other kava constituents,
lation leading to CDK1 activation in T24 cells. including four kavalactones of similar structures
Suppression of p53 expression by small interfer- to methysticin, demonstrated minimum activities
ing RNA in RT4 cells restored Cdc25C expres- in inhibiting NF-kappaB. A potent chalcone ana-
sion and down-regulated p21/WAF1 expression, log, (E)-3-(3-hydroxy-4-methoxyphenyl)-1-
which allowed Cdc25C and CDK1 activation, (3,4,5-trimethoxyphenyl)prop-2-en-1-one
which then led to a G(2)-M arrest and an enhanced derived from flavokawain compounds found in
growth-inhibitory effect by flavokawain kava, dose-dependently inhibited A549 lung can-
A. Consistently, flavokawain A also caused a pro- cer cell viability, NF-κB, activation of caspases
nounced CDK1 activation and G(2)-M arrest in and activation of mitogen-activated protein
p53 knockout but not in p53 wild-type HCT116 kinases such as extracellular signal regulated
cells. kinase 1/2 (ERK1/2) and c-Jun N-terminal kinase
Studies showed that a 30-week kava treatment (JNK) (Warmka et al. 2012)
inhibited 4-(methylnitrosamino)-1-(3-pyridyl)- Studies by Tang et al. (2010) demonstrated
1-butanone plus benzo[a]pyrene-induced lung that flavokawain B (FKB), a kava chalcone, was
tumorigenesis in A/J mice (Johnson et al. 2008). about 4- to 12-fold more effective in reducing the
Kava treatment inhibited proliferation and cell viabilities of androgen receptor (AR)-
enhanced apoptosis in lung tumors, as shown by negative, HRPC cell lines DU145 and PC-3 than
a reduction in proliferating cell nuclear antigen AR-positive, hormone-sensitive prostate cancer
(PCNA), an increase in caspase-3, and cleavage cell lines LAPC4 and LNCaP, with minimal
of poly(ADP-ribose) polymerase (PARP). Kava effect on normal prostatic epithelial and stromal
treatment also inhibited the activation of nuclear cells. FKB induced apoptosis with an associated
factor kappaBNF-kappaB, a potential upstream increased expression of proapoptotic proteins:
mechanism of kava chemoprevention. In subse- death receptor-5, Bim and Puma and a decreased
quent studies, Johnson et al. (2011) reported that expression of inhibitors of apoptosis protein:
mice-fed diets containing kava at dosages of XIAP and survivin. Additionally, FKB syner-
1.25, 2.5, 5 and 10 mg/g of diet had 8.4, 6.6, gized with TNF-related apoptosis-inducing
4.3 ± 2.4 and 3.8 lung adenomas per mouse, ligand (TRAIL) for markedly enhanced induc-
respectively corresponding to a reduction of tion of apoptosis. Furthermore, FKB treatment of
31 %, 46 %, 65 % and 69 % in tumor multiplicity. mice bearing DU145 xenograft tumors resulted
They found that flavokawains A, B and C, from in tumor growth inhibition and increased Bim
kava, demonstrated greatly reduced chemopre- expression in tumor tissues. Flavokawain B sig-
ventive efficacies even at concentrations much nificantly inhibited the cell proliferation of
higher than their natural abundance, suggesting human adenoid cystic carcinoma in-vitro in a
that they alone were unlikely to be responsible dose-dependent manner that was associated with
for kava’s chemopreventive activity. Kava at all induced apoptosis and cell cycle G2-M arrest,
dosages and treatment regimens did not induce and the half maximal inhibitory concentration
detectable adverse effects, particularly with (IC50) of flavokawain-B treatment for 48 oursh
respect to liver. Specifically, kava treatment was estimated to be 4.69 μmol/L (Zhao et al.
showed no effect on liver integrity indicator 2011). Induction of apoptosis was by up-
enzymes or liver weight, indicating that kava regulation of Bim and down-regulation of Bcl-2
may be potentially safe for long-term chemopre- expression. Li et al. (2012) treated prostate can-
ventive application. Shaik et al. (2009) demon- cer (LNCaP, LAPC-4, 22Rv1, C4-2B, DU145
strated that kava suppressed nuclear factor-kappaB and PC-3) cell lines having different androgen
(NF-kappaB) activation in lung adenoma tissues, receptor (AR) expression and a transformed
potentially a mechanism responsible for kava’s prostate myofibroblast cell line (WPMY-1), with
chemopreventive activity. Methysticin was iden- a commercial kava extract, kavalactones (kawain,
tified as a potent NF-kappaB inhibitor in kava 5′6′-dehydrokawain, yangonin, methysticin) and
Piper methysticum 157
flavokawain B. The kava extract and flavokawain tration of flavokawain A (FKA), the predominant
B effectively down-regulated the expression of chalcone from kava, to UPII-SV40T transgenic
both the full-length AR and AR splice variants. mouse for 318 days inhibited the occurrence of
The kava extract and kavalactones accelerated high-grade papillary urothelial cell carcinoma, a
AR protein degradation, while flavokawain B precursor to invasive urothelial cancer, by 42.1 %
inhibited AR mRNA transcription via decreasing (Liu et al. 2013). A decreased expression of Ki67,
Sp1 expression and the binding of Sp1 to the AR survivin and X-linked inhibitor of apoptotic pro-
promoter. The kava root extract and flavokawain teins (XIAP) and increased expression of p27 and
B reduced tumor growth, AR expression in tumor DR5, and the number of terminal deoxynucleoti-
tissues and levels of serum PSA in the patient- dyl transferase-mediated dUTP nick end labeling
derived prostate cancer xenograft in mice. The (TUNEL)-positive apoptotic cells were observed
results suggested a potential usefulness of a safe in the urothelial tissue of FKA-fed mice. The
kava product or its active components for preven- results suggested a potential of FKA in prevent-
tion and treatment of advanced prostate cancer by ing the recurrence and progression of non-
targeting AR. muscle-invasive urothelial cell carcinoma.
Flavokawain B (FKB) potently inhibited the The results of animal studies by Triolet et al.
growth of synovial sarcoma cell lines SYO-I and (2012) suggested that kava may reduce colon
HS-SY-II through induction of apoptosis (Sakai cancer risk. After 14 weeks, rats fed the nonpolar
et al. 2012). Treatment with FKB increased cas- fraction of kava extract 12 days prior to, during
pase 8, 9 and 3/7 activity compared to vehicle- and after the administration of dimethylhydra-
treated controls, indicating that both extrinsic and zine, a colon-specific carcinogen, had a signifi-
intrinsic apoptotic pathways were activated. In cant reduction in precancerous lesions [aberrant
addition, FKB treatment of both cell lines resulted crypt (AC) foci (ACF)] as well as large (≥ 4 AC/
in increased mRNA and protein expression of ACF) sialomucin-only expressing foci, an indica-
death receptor-5 and the mitochondrial pro- tor of greater tumorigenic potential, compared to
apoptotic proteins Bim and Puma, while down- the control group (Triolet et al. 2012). Groups fed
regulating the expression of an inhibitor of the ethanolic kava extract and polar kava fraction
apoptosis, survivin in a dose-dependent manner. trended toward reductions in ACF and large
Treatment with flavokawain B (FKB), a novel sialomucin-only expressing foci. The combined
kava chalcone, preferentially inhibited the growth kava groups had significantly fewer total AC,
of uterine leiomyosarcoma (SK-LMS-1), endo- ACF, large ACF and large sialomucin-only
metrial adenocarcinoma (ECC-1) cells compared expressing foci compared to the control group.
to the non-malignant, human endometrium Histological examination found no hepatic
fibroblast-like (T-HESC) cell lines (Eskander lesions in animals consuming the kava diets.
et al. 2012). FKB treatment resulted in cell cycle Studies by Leitzman et al. (2014) demonstrated
arrest and a robust induction of apoptosis in the outstanding efficacy of kava in preventing
SK-LMS-1 and ECC-1 cell lines. 4-(methylnitrosamino)-1-(3-pyridyl)-1-butanone
In a recent study, flavokawain B, a kava chal- (NNK)-induced lung tumorigenesis in A/J mice
cone, inhibited the growth of human osteosar- with high selectivity for the initiation stage in
coma cell lines 143B and Saos-2 through G2/M association with the reduction of O6-methylguanine
cell cycle arrest and induction of apoptosis adduct in DNA. Kava treatments covering the ini-
involving both extrinsic and intrinsic pathways tiation stage reduced the multiplicity of lung
(Ji et al. 2013). Furthermore, migration and inva- adenomas by approximately 99 %. A minimum
sion ability was decreased by FKB in a dose- effective dose was not defined because kava at
dependent manner. The cytotoxicity profile two lower dosages (2.5 and 1.25 mg/g of diet)
showed FKB had significant lower side effects on were equally effective as 5 mg/g of diet in
bone marrow cells and small intestinal epithelial completely inhibiting lung adenoma formation.
cells compared with adriamycin. Oral adminis- Dihydromethysticin from kava completely
158 Piperaceae
suppressed tobacco carcinogen 4-(methyl- Aurora B. Altogether, the results elucidated the
nitrosamino)-1-(3-pyridyl)-1-butanone-induced anti-inflammatory mechanisms triggered by tra-
lung tumorigenesis and differentially reduced ditionally used chemopreventive kava com-
DNA damage in A/J mice (Narayanapillai et al. pounds for inflammatory diseases and asthma.
2014a). Flavokawain A (FKA) from kava induced Of the kava-derived compounds tested in-
apoptosis in two breast cancer cell lines, MCF-7 vivo, kavain was found to render mice immune to
and MDA-MB231, and inhibited the metastatic lethal doses of lipopolysaccharide (Pollastri et al.
process in-vitro (Abu et al. 2014). FKA also halted 2009). Kavain demonstrated promising pharma-
the migration and invasion process in ceutical properties, including good solubility and
MDA-MB231. high cell permeability, but pharmacokinetic
experiments in mice showed relatively rapid
clearance. Of the kavain analogues synthesized, a
Antiinflammatory Activity ring-opened analog of kavain inhibited TNF-
alpha secretion in THP-1 cell-based assay and
Five known kawapyrones, 5,6-dehydrokawain suppressed lipopolysaccharide-induced TNF-
(isolated yield: 0.12 %), dihydrokawain (0.35 %), alpha factor expression in the same cells, whereas
kawain (0.47 %), yangonin (0.22 %) and methys- the other compounds inhibited TNF-alpha secre-
ticin (5.66 %) along with a new kavapyrone, tion without affecting lipopolysaccharide-
7,8-epoxyyangonin (6) (0.0059 %) isolated from induced TNF-alpha factor levels, indicating a
the methanol extract of dried kava powder were potential divergence in mechanism of action.
tested for TNF-α (tumor necrosis factor-α) Flavokawain A, from kava extract, significantly
release assay from BALB/3 T3 cells treated with suppressed expression of iNOS and COX-2, as
okadaic acid, a tumor promoter (Hashimoto et al. well as the subsequent production of NO and
2003). 5,6-Dehydrokawain (desmethoxyyango- PGE2 in the LPS-stimulated RAW 264.7 cells
nin) (1) and yangonin (4) significantly inhibited (Kwon et al. 2013). Flavokawain A significantly
TNF-α release with IC50 values of 17 μM and inhibited LPS-induced activation of NF-κB and
40 μM; a potency as great as (−)-epigallocatechin AP-1 signaling pathways. The results suggested
gallate (EGCG) isolated from green tea extract. that flavokawain A may exert anti-inflammatory
Among the kavapyrones, dihydrokawain was responses by suppressing LPS-induced expres-
unique in showing the strongest inhibitory activ- sion of pro-inflammatory mediators via blockage
ity against TNF-α release in mice, but the weak- of NF-κB-AP-1-JNK/p38 MAPK signaling path-
est activity in the cells. Studies by Folmer et al. ways in the murine macrophages. Studies showed
(2006) found that 320 μM (flavokavain A), that a kavalactone derivative2′,6′-dichloro-5-
175 μM (flavokavain B) and 870 μM (kavain and methoxymethyl-5,6-dehydrokawain inhibited
dihydrokavain) inhibited both NF-kappa B-driven lipopolysaccharide-stimulated iNOS induction
reporter gene expression and TNF alpha-induced and NO production through activation of Nrf2
binding of NF-kappaB. Moreover, kavain and fla- signaling in BV2 microglial cells (Terazawa et al.
vokavains A and B treatment led to inhibition of 2013). The data suggested that the compound had
both inhibitors of kappaB (IkappaB) degradation a potential to reduce neuroinflammation as well
and subsequent translocation of p50 and p65 as oxidative stress in neurodegenerative diseases
NF-kappa B subunits from the cytoplasm to the through activation of Nrf2 signaling.
nucleus. Further, kinase selectivity screening
demonstrated that flavokavain A, but not kavain,
nor flavokavain B, inhibited the IkappaB kinase Central Nervous Sytem (CNS) Activity
(IKK) as well as PRAK (p38-regulated/activated
kinase), MAPKAP-K3 (MAPK-activated protein Animal studies showed that kava lactones altered
kinase 3), DYRK1A (dual-specificity tyrosine- neuronal excitation through direct interactions
phosporylated and regulated kinase 1A) and with voltage-dependent ion channels, giving rise
Piper methysticum 159
tion of the anxiolytic chlordiazepoxide (CDP) by nificantly reduced anxiety symptoms after
kava extracts. Partial substitution of kava extract 1 week; no adverse effects were noted during the
for CDP suggested that kava may share a mecha- 4-week treatment (Kinzler et al. 1991). In a one
nism of action similar to CDP, but was less 6-month, double-blind study compared kava was
potent. found to be just as effective as two standard anxi-
ety benzodiazepine drugs (oxazepam and bro-
Clinical Studies mazepam) in 174 people with anxiety symptoms
Two placebo-controlled trials investigated the (Woelk et al. 1993). Improvement in Hamilton
effect of standardized Kava extracts in women Anxiety Scale (HAMA) scores was about the
with climacteric psychosomatic disturbances same in all groups. A 4 weeks, randomized,
(Warnecke et al. 1990; Warnecke 1991). In the placebo-controlled double-blind study of 58
first study involving 40 such women, kava extract patients also assessed the efficacy of WS 1490
(200–400 mg extract = 30–60 mg kava pyrones) extract containing 70 % kava pyrones for the
for 8 weeks was found to be superior to the pla- treatment of anxiety of non-psychotic origin
cebo using the Kuppermann Index and Anxiety (Lehmann et al. 1996). The Hamilton Anxiety
Status Index. In the second randomized, placebo- Scale (HAMA) overall score of anxiety symp-
controlled double-blind study, two groups each tomatology revealed a significant reduction in the
containing 20 patients with climacteric-related kava group compared with the placebo group
symptomatology were treated for a period of 8 after 1 week of treatment. The results of the sec-
weeks with kava WS 1490 extract 3 × 100 mg/day ondary target variables were in agreement with
or a placebo preparation. The Hamilton Anxiety the HAMA score and demonstrated the efficacy
Scale (HAMA) overall score of anxiety symp- of WS 1490 in patients with anxiety disorders. In
tomatology revealed a significant difference in another randomized placebo-controlled 25-week
the kava-receiving group compared to the pla- outpatient trial of 101 outpatients suffering from
cebo group already after only 1 week of treat- anxiety of non-psychotic origin (Diagnostic and
ment. The course of such further parameters as Statistical Manual of Mental Disorders (DSM)
depressive mood (DSI), subjective well-being -III-R criteria: agoraphobia, specific phobia, gen-
(patient diary), severity of the disease (CGI) and eralized anxiety disorder and adjustment disorder
the climacteric symptomatology (Kuppermann with anxiety), treatment with kava-kava extract
Index and Schneider scale) over the overall WS 1490 was found to be superior over placebo
period of treatment demonstrated a high level of from week 8 onwards in the Hamilton Anxiety
efficacy of kava extract WS 1490 in neurovegeta- Scale (HAMA) (Volz and Kieser 1997). WS 1490
tive and psychosomatic dysfunctions in the cli- was also found to be superior with respect to the
macteric, associated with very good tolerance of secondary outcome variables. The results sup-
the preparation. A randomized, double-blind ported WS 1490 as a treatment alternative to tri-
pilot study investigated the effects of the extract cyclic antidepressants and benzodiazepines in
containing 15 % kava pyrones in 59 patients with anxiety disorders. In an outpatient observation
pre-operative anxiety (Bhate et al. 1989). study of 52 outpatients suffering from anxiety of
Although improvements in mood were observed nonpsychotic origin, the treatment of kava extract
using a psychostatus score, only two doses of the was found to be as an effective and safe alterna-
extract (equivalent to 60 mg kava pyrones daily) tive to antidepressants and tranquilizers in anxi-
were administered, and thus the clinical signifi- ety disorder without the tolerance problems
cance of this study was questionable. associated with benzodiazepines (Scherer 1998).
In a placebo-controlled double-blind study of The results of a randomised study of 40 women
29 patients with anxiety syndromes, treatment in physiological or surgical menopause for the
with 100 mg of WS 1490 three times a day sig- past 1–12 years showed that the association of
162 Piperaceae
hormone replacement therapy (HRT) and kava- the same extract. WS 1490 was well tolerated,
kava extract may represent an excellent therapeu- with no influence on liver function tests and only
tic tool for the treatment of women in stabilized one trivial adverse event (tiredness) attributable
menopause, in particular those suffering from to the study drug. In a 3-week placebo-controlled,
anxiety and depression, given that kava-kava double-blind crossover trial that recruited 60
therapy accelerated the resolution of psychologi- adult participants with 1 month or more of ele-
cal symptoms without diminishing the therapeu- vated generalized anxiety, administration of 5
tic action of estrogens on organic disease, such as kava tablets per day containing 250 mg of kava-
osteoporosis and cardiovascular disease (De Leo lactones/day, produced significant anxiolytic and
et al. 2000). antidepressant activity and raised no safety con-
Malsch and Kieser (2001) conducted a 5-week cerns at the dose and duration studied (Sarris
randomised, placebo-controlled, double-blind et al. 2009b). Kava appeared equally effective in
study to investigate the efficacy of kava-kava spe- cases where anxiety was accompanied by depres-
cial extract WS1490 in 40 patients with non- sion. In another 3-week placebo-controlled,
psychotic nervous anxiety, tension and double-blind, cross-over trial involving 60 adult
restlessness states. The study confirmed the anx- participants (18–65) with elevated stable anxiety
iolytic efficacy and good tolerance of WS1490 and varying levels of depressive symptoms, kava
over placebo and showed that a further symptom preparation exerted significant anxiolytic effects
reduction was possible after a change over from and appeared safe (Sarris et al. 2009a). In a ran-
benzodiazepine treatment. Studies by Watkins domized, placebo-controlled, double-blind study
et al. (2001) suggested that kava might exert a of 22 moderately anxious adults aged between 18
favourable effect on reflex vagal control of heart and 65 years, kava administration was found to
rate in generalized anxiety disorder patients. have no negative effect on cognition, whereas a
Significantly more patients treated with kava reduction in alertness occurred in the oxazepam
showed improved baroreflex control of heart rate condition (Sarris et al. 2012). Genetic analyses
(BRC) compared to the placebo group. Further, provided tentative evidence that noradrenaline
the magnitude of improvement in BRC was sig- (SLC6A2) transporter polymorphisms may have
nificantly correlated with the degree of clinical an effect on response to kava. They concluded
improvement. In a 3-month randomized prospec- that acute “medicinal level” doses of the particu-
tive open study of perimenopausal women, the lar kava cultivar in naive users did not provide
administration of kava-kava plus calcium elicited anxiolytic activity, although the phytomedicine
an improvement of mood, particularly of anxiety also appeared to have no negative effects on
(Cagnacci et al. 2003). Gastpar and Klimm cognition.
(2003) investigated the efficacy and tolerability In a more recent 6-week double-blind, ran-
of 150 mg/day kava special extract WS 1490 ver- domized, placebo-controlled study of 75 partici-
sus placebo in a randomized, placebo-controlled, pants with generalized anxiety disorder (GAD)
double-blind multicenter study in 41 adult, male and no comorbid mood disorder, treatment with
and female out-patients suffering from neurotic kava produced a significant reduction in anxiety
anxiety wherein patients received 3 × 1 capsule of compared with the placebo group with a moder-
50 mg/day WS 1490 or placebo for 4 weeks, fol- ate effect size (Sarris et al. 2013b). Among par-
lowed by 2 weeks of observation without study- ticipants with moderate to severe Diagnostic and
specific treatment. The results showed consistent Statistical Manual of Mental Disorders-diagnosed
advantages for WS 1490 over placebo in several GAD, this effect was larger. The data suggested
psychiatric scales and indicated significant that standardized kava may be a moderately
improvements in the patients’ general well-being, effective short-term option for the treatment of
the differences versus placebo were not as large GAD. Furthermore, specific γ-aminobutyric acid
as in previous trials which employed 300 mg/d of (GABA) transporter polymorphisms appeared to
Piper methysticum 163
potentially modify anxiolytic response to kava. 1959; Meyer 1962; Meyer and May 1964). Frater
In another 6-week, double-blind, randomized (1958) demonstrated that a thin paste of kava
controlled trial (n = 75) involving chronic admin- powder applied to the lip mucous membrane pro-
istration of kava or placebo for participants with duced a slightly burning sensation and a feeling
generalized anxiety disorder, no significant dif- of numbness when some kava roots were chewed
ferences were found across groups for liver func- for 15 min found the degree of anaesthesia was
tion tests nor were there any significant adverse greater. Steinmetz (1960) described the narcotic
reactions that could be attributed to kava (Sarris effects of kava as a paralysis of the nervous sys-
et al. 2013c). Interesting, kava significantly tem, through reduction of spinal rather than cere-
increased female’s sexual drive compared to pla- bral activity, followed by muscular stimulation
cebo on a sub-domain of the Arizona Sexual and then paralysis particularly affecting the lower
Experience Scale (ASEX), with no negative limbs. Also noted was a reduction of the cardiac
effects seen in males. Further, it was found that rhythm followed by stimulation and slowing
there was a highly significant correlation between down of respiration. Steinmetz also stated that
ASEX reduction (improved sexual function and ingestion of a strong dose would affect eye vision,
performance) and anxiety reduction in the whole causing pupil dilation and inducing photophobia.
sample. A gel containing 6 % kava root crude extract
called kavacaine was found to have potent topical
Meta-analysis/Review Studies anaesthetic effect (Morse and Sharma 2005).
In a meta-analysis of 11 randomised, controlled Administration of dihydromethysticin to mice
trials (RCTs), i.e. trials with a randomised gen- potentiated pentobarbital-induced sleeping time
eration of allocation sequences, and conducted by 400 %, while dihydrokawain, yangonin and
placebo-controlled and double-blind, i.e. trials kawain were only moderately active (150–235 %)
with blinding of patients and care providers and (Klohs et al. 1959). Both dihydrokawain and
involving a total of 645 participants, Pittler and dihydromethysticin exhibited analgesic effects
Ernst (2003) found that kava extract appeared to when administered intraperitoneally to rats
be an effective symptomatic treatment option for (140 mg/kg body weight), as determined by an
anxiety compared to placebo. The data available increase in tail-flick reaction times (Brüggemann
from the reviewed studies suggested kava to be and Meyer 1962). Meyer (1962) found that
relatively safe for short-term treatment (1–24 administration of 150 mg/kg of hexobarbital
weeks), although more information was required. sodium to white mice afforded 2 h of sleep.
In a comprehensive review of efficacy, safety and However, pretreament with 240 mg/kg dihydro-
psychopharmacology, Sarris et al. (2011) found methysticin induced 72 h of sleep. Meyer and
that current evidence supported the use of kava in May (1964) found all the kava lactones acted as
treatment of anxiety with a significant result local analgesics. Most of the kava lactones inhib-
occurring in four out of six studies reviewed. It ited frog heart contraction. These actions were
was suggested that more studies were required to comparable to cocaine which also protected
assess comparative efficacy and safety (on the against ventricular fibrillation through its local
liver, cognition, driving and sexual effects) ver- anaesthetic effect. Dihydroymethysticin and
sus established pharmaceutical comparators. dihydrokavain had been shown to intensify sleep-
inducing effects (Hänsel 1968). Both compounds
were found to be comparable to the drug dimeth-
Sedative/Analgesic/Anaesthetic/ ylaminophenazone in producing an analgesic
Anti-insomnia Activities effect in the drinker. The analgesic effect was
found to be stronger than aspirin but weaker then
The local anaesthetic activity of kava was first morphine. Kavain had been shown to produce
observed by Lewin (1886). Kava had been local aesthetic effect that was comparable to
reported to possess and to potentiate and prolong cocaine and do not produce any toxicity in the
barbiturate narcosis (Frater 1958; Klohs et al. tissues (Kretzschmar and Meyer 1969).
164 Piperaceae
The extent of frog sartorius muscle paralysis been reported to be taken in conjunction with
induced by kava was similar in both directly and alcoholic drinks. Psychopharmacological effects
indirectly stimulated mouse hemidiaphragms of kava had also been reported and the mood-
(Singh 1983). Intracellular recordings from frog altering effects of kavalactones had been
sartorius muscles showed that kava depressed the described as hypnotic (Schultes and Hoffman
amplitude of both miniature end-plate potentials 1992). The pharmacological properties of kava-
(mepps) and end-plate potentials (epps) but had lactones were comparable to those of benzodiaz-
no effect on the frequency of mepps. Kava greatly epines, however, kavalactones bind very weakly
prolonged the duration of mepps and epps and to the gamma-aminobutyric acid (GABA) and
also slowed and depressed directly elicited mus- benzodiazepine (Bilia et al. 2002). It was pro-
cle action potentials. The neuromuscular block- posed that N-methyl-d-aspartate receptors and/or
ade produced was poorly reversed by calcium voltage dependent calcium channels may also be
and by neostigmine. It was concluded that kava involved in the elementary mechanism of action.
caused paralysis by mechanisms similar to local Only weak activity of kava resin was observed on
anaesthetics. Both the intoxicating beverage kava GABAA binding sites washed synaptosomal
and the lipid soluble (kava resin) extracts showed membranes prepared from rat brain and this was
analgesic effects in both tail immersion and abolished by extraction of the membranes with
abdominal constriction in mice (Jamieson and Triton X-100, suggesting that lipid soluble com-
Duffield 1990b). Eight purified pyrones from the ponents were involved (Davies et al. 1992). No
lipid soluble extract were also tested for activity effects were observed on GABAB binding sites
in the tail immersion test, and kawain, dihydro- in rat brain membranes in-vitro. Kava resin and
kawain, methysticin and dihydromethysticin pyrones exerted some weak effects on benzodiaz-
were found to be very effective in producing epine binding in-vitro but this did not correlate
analgesia. Naloxone, in doses which inhibited with pharmacological activity. In addition, in ex-
morphine-induced analgesia in both tests, was vivo studies, no effects were observed on [3H]
indicating that analgesia produced by kava diazepam binding to brain membranes prepared
occurred via non-opiate pathways. The synthe- from mice in which selected kava constituents
sized analogues of synthetic kavain, were injected intraperitoneally, whereas similarly
6-(4-fluorophenyl)-4-methoxy-5,6-dihydropyran- administered diazepam (5 mg/kg) inhibited [3H]
2-one exerted potent and dose-dependent diazepam binding by greater than 95 %. Similar
analgesic activity, inhibiting abdominal constric- lack of activity was observed in in-vivo binding
tions caused by acetic acid in mice, and being studies; injection of kava resin failed to influence
more active than some reference drugs (Kormann the CNS binding of the benzodiazepine-receptor
et al. 2012). It also presented activity in the other ligand [3H]Ro15-1788 injected into mice prior to
models of pain, with the exception of the hot sacrifice. The pharmacological activities of kava
plate test and the measurement of motor resin and pyrones did not appear to be elucidated
performance. by any significant interaction with GABA or ben-
The lipid soluble extract of the psychoactive zodiazepine binding sites. Studies by Jussofie
beverage kava was found to have hypnosedative et al. (1994) indicated that one way kavapyrone-
properties which could be measured by the length enriched kava extract might mediate sedative
of time that the righting reflex was lost (Jamieson effects in-vivo was through effects on gamma-
and Duffield 1990a). Ethanol and the lipid solu- aminobutyric acid (GABA) receptor binding.
ble extract (kava resin) was shown greatly to The sedative effect was demonstrated using
increase each others’ hypnotic action in mice. membrane fractions obtained from target brain
Ethanol also increased the toxicity of kava mark- centers of kavapyrone action: hippocampus
edly. This interaction of kava and alcohol pos- (HIP), amygdala (AMY) and medulla oblongata
sessed important clinical and social consequences (MED), and from brain centers outside the main
since, in contrast to traditional usage, kava had kavapyrone effects as frontal cortex (FC) and cer-
Piper methysticum 165
ebellum (CER). At a kavapyrone concentration of do not produce any toxicity in the tissues
500 μM, the order of enhancement in binding (Kretzschmar and Meyer 1969). Examination of
sites (Bmax) was HIP = AMY > MED > FC > CE EEG recordings during sleep of healthy volun-
R. When kavapyrones were included together teers given a single dose of 300 mg extract
with pentobarbital or 3-alpha-hydroxy-5-alpha-- (equivalent to 210 mg kava pyrones) showed an
pregnane-20-one (HPO), the two classes of com- increased sleep spindle density of 20 % and an
pounds produced a synergetic effect on [3H] increase in slow-wave sleep (i.e. deep sleep), but
muscimol binding. the rapid eye movement phase was not sup-
In a double-blind, placebo-controlled study, pressed (Emser and Bartylla 1991). Daily doses
the encephalotropic and psychotropic effects of of 300 or 600 mg extract (equivalent to 210 or
kavain-a synthetic kava plant derivative as com- 420 mg kava pyrones), respectively, for 1 week
pared with clobazam were investigated in 15 nor- increased the beta/alpha index typical for the
mal volunteers, utilizing EEG brain mapping, pharmaco-EEG profile of anxiolytics without the
psychometric and psychopysiological analyses sedative-hypnotic effects associated with benzo-
(Saletu et al. 1989). Brain maps of drug induced diazepines. The increase in beta activity was
pharmaco-EEG changes (pharmaco-EEG maps) most marked in the beta2 range (Johnson et al.
demonstrated that kavain exerted a significant 1991). In two studies, administration of 300 mg
action on the human brain function as compared kava extract WS 1490 (equivalent to 210 mg kava
with placebo characterized by a dose-dependent pyrones) daily for either 8 or 14 days to healthy
increase of delta, theta and alpha 1 activity while volunteers, taken with or without ethanol, had no
alpha 2, beta activity and the centroid of the total potentiating or additive effect and had no influ-
activity decreased. These findings were indica- ence on the safety-related performance of healthy
tive of a sedative effect which was, however, in volunteers (Herberg 1991, 1993). In another
type quite different from that of the study administration of kava extract standardized
1,5-benzodiazepine. Further, 200 mg kavain to contain 30 % kava pyrones, bromazepam or a
induced with a decrease of delta and beta activity combination of extract and bromazepam, safety-
and an increase of alpha activity and of total related performance remained unaffected in
power also vigilance promoting effects. Kavain healthy volunteers treated daily with 400 mg
improved the noopsyche as compared with pla- extract (equivalent to 120 mg kava pyrones for 14
cebo in all three doses as there was a significant days), whereas it was impaired after treatment
improvement in intellectual performance (Pauli with bromazepam (9 mg daily) or the extract/bro-
test), attention, concentration, reaction time and mazepam combination (Herberg 1996). No dif-
motor speed (rigidity test), while opposite find- ferences were observed following treatment with
ings were observed after 30 mg clobazam. In bromazepam or the combination, indicating that
regard to thymopsychic variables such as drive, the extract did not have an additive effect when
wakefulness, affectivity, mood and well-being, given in combination with bromazepam. In a
200 mg kavain produced an improvement as separate, randomized, double-blind crossover
compared with placebo while 600 mg kavain pro- study involving 12 healthy volunteers, adminis-
duced sedation as did 30 mg clobazam. tration of daily single doses of a kava extract
Dihydroymethysticin and dihydrokavain had standardized to contain 30 % kava pyrones
been shown to intensify sleep-inducing effects (400 mg extract containing 120 mg kava pyrones)
(Hänsel 1968). Both compounds were found to was compared with daily single doses of diaze-
be comparable to the drug dimethylaminophena- pam (10 mg) or a placebo in a 7-day trial (Gessner
zone in producing an analgesic effect in the and Cnota 1994). Changes in EEG recordings
drinker. The analgesic effect was found to be and psychometric test results showed no evidence
stronger than aspirin but weaker than morphine. of a decrease in vigilance in the group treated
Kavain had been shown to produce local aes- with the extract. Following administration of
thetic effect that was comparable to cocaine and kava-kava extract at a dose of 300 mg/kg, a sig-
166 Piperaceae
nificant shortening of the sleep latency in sleep- the lipid-soluble extract (kava resin) reduced
disturbed rats was observed while no effects were amphetamine-induced hypermotility (Duffield
observed on the total waking and non-rapid eye et al. 1989b). Aqueous kava extract in i.p. doses
movement (non-REM) sleep time (Shinomiya of 30 mg/kg to 500 mg/kg had no effect on con-
et al. 2005). In contrast, flunitrazepam showed a ditioned avoidance responses. At or below
significant shortening in sleep latency, decrease 100 mg/kg i.p., kava resin also failed to modify
in total waking time and increase in total non- the number of conditioned avoidance responses
REM sleep time. Although the effects of fluni- obtained. However, 125 mg/kg of resin signifi-
trazepam were antagonized by the benzodiazepine cantly reduced the number of conditioned avoid-
receptor antagonist flumazenil, the effect of kava- ance responses by 18 %. Increasing the dose of
kava extract was not antagonized by flumazenil. kava to 150 mg/kg caused ataxia and sedation. A
Kava-kava extract showed a significant increase minimally effective daily dose (50 mg/kg) of the
in delta activity during non-REM sleep in sleep- aqueous kava extract for 3 days was sufficient to
disturbed rats, whereas a significant decrease in produce tolerance to a test dose of 150 mg/kg,
delta power during non-REM sleep was observed which was close to the ED50 (Duffield and
with flunitrazepam. The results showed that Jamieson 1991). Kava resin decreased spontane-
Kava-kava had not only hypnotic effects but also ous motility and caused a loss of muscle control.
sleep quality-enhancement effects. A minimally effective daily dose of kava resin
Two fractions, F1 and F2, from kava extract (100 mg/kg) did not produce tolerance to the
were found to decrease spontaneous motor activ- above effects of a weekly test dose of kava resin
ity in doses which did not alter forced motor (166 mg/kg) within 7 weeks. Increasing the dose
activity of mice (O’Hara et al. 1965). Fraction to 150 mg/kg twice daily caused partial tolerance
F2, dihydromethysticin, desmethoxy-yangonin to occur within 3 weeks, but very little further
and kawain exhibited potent antiserotonin activ- tolerance developed over the ensuing 2-week
ity on the isolated rat uterus, whereas F1 appeared period. Intraperitoneal administration of an
to be devoid of antiserotonin activity. F1, F2 and extract of kava rhizome (equivalent to 50–100 mg
dihydromethysticin did not alter serotonin brain kava pyrones/kg body weight) or (±)-kawain, a
levels in mice. Intraperitoneal (i.p.) adminstra- synthetic kava lactone (10–50 mg/kg body
tion of an aqueous extract of kava (62.5 mg/kg weight), reduced muscle tone in cats (Holm et al.
body weight) caused a loss of spontaneous activ- 1991). Intraperitoneal administration of kava rhi-
ity without loss of muscle tone in mice (Jamieson zome (equivalent to 50–100 mg kava pyrones/kg
et al. 1989). No hypnotic effect was seen, but body weight) to cats had a significant effect on
some analgesia was produced. The anticonvul- EEG recordings, inducing high-amplitude delta
sant effect against strychnine was very slight and waves, spindlelike formation and continuous
there was no evidence of local anesthetic action. alpha- or beta-synchronization in amygdala
There was a slight anti-apomorphine effect and recordings. Hippocampal responses, following
tetrabenazine-induced ptosis was decreased. The stimulation of the amygdala nucleus, increased
lipid-soluble extract (kava resin) also decreased significantly in amplitude in cats treated intra-
spontaneous motility and markedly reduced peritoneally with the rhizome extract or
motor control (Jamieson et al. 1989). Hypnosis, (±)-kawain.
determined by loss of righting reflex, was pro-
duced, analgesia was marked, and a local anes-
thetic action was evident. Kava resin also Anticonvulsant Activity
decreased apomorphine-induced hyper-reactivity
and partially reversed tetrabenazine-induced pto- Intraperitoneal administration of an aqueous
sis in rats. Kava resin produced a greater range of extract (300 mg/kg body weight) or a chloroform
pharmacological actions than the aqueous extract, extract (140 mg/kg body weight) of the rhizome
and the latter was orally inactive in mice and rats. to mice-inhibited strychnine-induced convul-
The aqueous, pyrone-free extract from kava and sions (Klohs et al. 1959). The anticonvulsant
Piper methysticum 167
activity of methysticin and other kava pyrones slices containing the hippocampus and the ento-
against electroshock- and chemically induced rhinal cortex, methysticin in a concentration
seizures has been demonstrated in mice and rats range of 10–100 μM reversibly blocked all types
(Keller and Klohs 1963; Meyer and Meyer-Burg of epileptiform activity (Schmitz et al. 1995).
1964; Kretzschmar and Meyer 1969; Kretzschmar While responses to alvear stimulation were
et al. 1970). Intraperitoneal administration of largely unaffected by methysticin, the responses
dihydromethysticin and dihydrokawain inhibited to a paired pulse stimulus to stratum radiatum
electroshock-induced seizures at doses of 25 and were depressed over the whole range of tested
60 mg/kg body weight, respectively, in mice stimulus intervals. The findings suggested that
and rats (Meyer and Meyer-Burg 1964). methysticin had effects on different patterns of
Dihydrokavain and dihydromethysticin inhibited epileptiform activity possibly by interfering with
muscular contractions and the effect was found to processes responsible for frequency potentiation.
be comparable to those of synthetic products Both (±)-kawain and methysticin inhibited
such as phenobarbital (Kretzschmar and Meyer voltage-dependent sodium channels in rat CA1
1969). Dihydromethysticin was reported to have hippocampal neurons in-vitro (1–400 μmol/l)
strong anticonvulsant action with the capability (Magura et al. 1997). In cultured dorsal root gan-
of inhibiting convulsions caused by strychnine in glion cells derived from neonatal rats,
animals (Kretzschmar et al. 1970). Both Schirrmacher et al. (1999) found that (+/−)-kavain
compounds were reportedly more superior to reduced currents through voltage-activated Na+
benzodiazepine as muscle relaxants. It had and Ca2+ channels.
been suggested that the anti-epileptic action of Kava pyrones were reported to exert effects on
dihydryomethysticin may be used to treat schizo- neuronal transmission and transmembraneous
phrenia. Isolated kavain, methysticin, dihydro- cation currents similar to established mood stabi-
methysticin and yangonin showed strong lizers like carbamazepine, valproate and
centrally mediated muscle relaxing activity in lamotrigine (Grunze et al. 2001). Findings of
rabbits and yangonin was the most potent kava- studies suggested that (i) kava pyrones had a
lactone (Kretzschmar et al. 1971). Studies had weak Na+ antagonistic effect that may contribute
shown kava pyrones to be anticonvulsive (Gleitz to their antiepileptic properties; (ii) they had pro-
et al. 1996b). The anticonvulsive action of nounced l-type Ca2+ channel antagonistic proper-
(+/−)-kavain was estimated from its properties on ties and acted as a positive modulator of the early
stimulated synaptosomes and Na+ channel recep- K+ outward current; two actions of importance
tor sites. The results suggested an interaction of for mood stabilization; (iii) kava pyrones had
(+/−)-kavain with voltage-dependent Na+ and additive effects with the serotonin-1A agonist
Ca2+ channels, thereby suppressing the ipsapirone probably contributing to their anxio-
4-aminopyridine-induced increase in [Na+]i, lytic and sleep-inducing effects; and (iv) kava
[Ca2+]i and the release of endogenous glutamate. pyrones showed a distinct pattern of action on
(+/−)-Kavain specifically and rapidly inhibited glutamatergic and GABAergic transmission
veratridine-activated voltage-dependent Na+- without affecting long-term potentiation. It was
channels in synaptosomes prepared from rat found that kava pyrones exhibited a profile of cel-
cerebral cortex (Gleitz et al. 1995). Further lular actions that showed a large overlap with
they found that (+/−)-kavain inhibited the several mood stabilizers, especially lamotrigine.
veratridine-induced and KCl-induced increase in
intracellular Ca2+ and glutamate-release of rat
cerebrocortical synaptosomes (Gleitz et al. Neuroprotective Activity
1996a). The results suggested that (+/−)-kavain
at concentrations sufficient to block Na+ channels The neuroprotective effects of an acetone extract
completely, moderately inhibited the non- of kava rhizome and kava pyrones had been dem-
inactivating Ca2+ channels located on mammalian onstrated both in-vivo and in-vitro. Administration
presynaptic nerve endings. In rat temporal cortex of kava extract (orally) and its constituents
168 Piperaceae
methysticin and dihydromethysticin (intraperito- effects of oxazepam and kava root extract on
neally) protected against brain tissue against behavior and event-related potentials (ERPs) in
ischemic damage in mice and rats (Backhauss a recognition memory task in twelve healthy
and Krieglstein 1992). The kava extract (150 mg/ volunteers. Oxazepam led to a reduction of a
kg, 1 h before ischemia) diminished the infarct negative component in the 250–500 ms range
area in mouse brains and the infarct volume in rat for both old and new words and to a reduction of
brains. Methysticin, dihydromethysticin (both 10 the old/new difference in the ERP associated
and 30 mg/kg, 15 min before ischemia) and the with a significantly worse recognition rate.
reference substance memantine (20 mg/kg, Kava, on the other hand, showed a slightly
30 min before ischemia) significantly reduced the increased recognition rate and a larger ERP dif-
infarct area in mouse brains. The standardized ference between old and new words Heinze
extract also protected against neuronal damage in et al. (1994) investigated the effects of oxaze-
cultured neurons from chick embryo cerebral pam and a standardized extract of kava roots
hemispheres (Backhauss and Krieglstein 1992). (WS1490) on reaction time and event-related
Kava pyrones exhibited neuroprotective and potentials (ERPs) in a visual search paradigm
“recovery-supporting” effects on neurological using a double-blind design in young, healthy
deficits after cerebral infarction in rats (Kleiser males. Significant effects of oxazepam were
et al. 1998). obtained in a number of psychometric tests.
Administration of the lower dosages of Oxazepam led to a reduction of the amplitude of
(+/−)-kavain (50 and 100 mg/kg) to male the parietal N1, frontal N2, posterior contralat-
C57BL/6 mice afforded only a nonsignificant eral N2 and occipital P3 components. WS 1490
attenuation of MPTP (1-methyl-4-phenyl- was associated with a greater posterior N1, pos-
1,2,3,6-tetrahydropyridine)-induced dopamine terior contralateral N2 and occipital P3. In a
depletion, but a high dosage of (+/−)-kavain double-blind randomized placebo-controlled
(200 mg/kg) significantly antagonized the dopa- trial involving healthy volunteers, Thompson
mine depletion to 58.93 % of saline control val- et al. (2004) found the intake of a single dose of
ues (Schmidt and Ferger 2001). The kava extract (300 mg; p.o.) led to an increase in
MPTP-induced decrease of tyrosine hydroxylase- state cheerfulness, while the herbal medicine
immunoreactivity as well as the loss of nigral did not influence state seriousness and bad
neurons was completely prevented by mood. The mood-elevating effects of kava were
(+/−)-kavain (200 mg/kg). They concluded that most prominent in trait cheerful subjects, indi-
MPTP metabolism was not influenced by cating that trait cheerfulness moderated the
(+/−)-kavain and postulated the antiglutamater- drug-induced increase in cheerful mood.
gic effects of (+/−)-kavain for its protective Further, Kava improved the accuracy and the
effects against MPTP toxicity. They asserted that speed of performing the partial report and the
kavain may be a novel candidate for further pre- item recognition task, indicative of a beneficial
clinical studies in animal models of Parkinson’s effect of kava on visual attention and short-term
disease and other disorders with glutamatergic memory retrieval, respectively.
overactivity. In a systematic review of clinical trials on the
effects of kava on cognition, La Porte et al. (2011)
found the majority of evidence from 10 human
Cognitive Activity clinical trials suggested that kava had no repli-
cated significant negative effects on cognition.
Kava was found to have no effect on the reaction One acute study found that kava significantly
times or errors of two groups of undergraduates improved visual attention and working memory
who consumed doses thought usual (Experiment processes while another found that kava increased
1) and doses much greater (Experiment 2) than body sway. One chronic study found that kava
those associated with social functions (Russell significantly impaired visual attention during
et al. 1987). Münte et al. (1993) investigated the high-cognitive demand.
Piper methysticum 169
Antimicrobial Activity
Melanogenesis Stimulation Activity
Kavain was reported to exhibit bactericidal activ-
P. methysticum rhizome ethanol extract showed ity against Gonococcus, the pathogenic agent of
potent stimulatory effect on melanogenesis in gonorrhoea and colon bacillus and blennorrhea
cultured murine B16 melanoma cells (Matsuda (Steinmetz 1960). The pathogenic fungi
et al. 2006). Activity-guided fractionation of Trichophyton ferrugineum, Trichophyton
kava extract led to the isolation of two active kav- tonsurans and Cryptococcus neoformans were
alactones, yangonin and 7,8-epoxyyangonin, strongly inhibited by kava extracts (Hänsel et al.
along with three inactive kavalactones, 1966a). Dihydromethysticin completely inhib-
5,6-dehydrokawain, (+)-kawain and (+)-methys- ited the growth of Aspergillus niger in-vitro
ticin, and a glucosylsterol, daucosterin. (Shulgin 1973). The fungistatic principles of the
7,8-Epoxyyangonin showed a significant stimu- kava root were found to be 4-methoxy-α-pyrones
latory effect on melanogenesis in B16 melanoma like dihydrokawain which completely inhibited
cells. Yangonin exhibited a weak melanogenesis Aspergillus niger at a concentration of 0.5 mg/
stimulation activity. ml. Other fungi (species of Candida and
Fusarium) and bacteria were not inhibited by
kava and dihydrokawain. A hydroalcoholic
Antiplatelet Activity extract of kava rhizome inhibited the growth in-
vitro of Aspergillus fumigatus, A. niger,
Kava-kava extract was found to be a reversible Penicillium digitatum, Rhizopus nigricans,
inhibitor of MAO-B in intact platelets (IC50 Trichophyton mentagrophytes, Candida albicans
24 μM) and disrupted platelet homogenates (IC50 and Saccharomyces pastorianus (Guérin and
1.2 μM) (Uebelhack et al. 1998). The order of Réveillère 1984). However, an aqueous extract of
potency was desmethoxyyangonin > (±)-methys- the rhizome weakly inhibited the growth in-vitro
ticin > yangonin > (±)-dihydromethysticin > (±)- of Trichophyton rubrum, Microsporum canis or
dihydrokavain > (±)-kavain. The two most potent Epidermophyton floccosum (Locher et al. 1995).
Piper methysticum 171
Treatment of F-344 rats with pipermethystine methysticin triggered the most profound induc-
(10 mg/kg) and kava root acetone-water extract ing effect on CYP1A1 followed by 7,8-dihydro-
(100 mg/kg) for 2 weeks failed to elicit any sig- methysticin. The other four kavalactones
nificant changes in liver function tests or cause (yangonin, 5,6-dehydrokawain, kawain and
severe hepatic toxicity as measured by lipid per- 7,8-dihydrokawain) did not show significant
oxidation and apoptosis markers such as malo- effects on CYP1A1. It was found that kava
ndialdehyde, Bax and Bcl-2 (Lim et al. 2007). extract induced the expression of CYP1A1 via an
However, pipermethystine-treated rats demon- aryl hydrocarbon receptor (AhR)-dependent
strated a significant increase in hepatic glutathi- mechanism and that methysticin and
one, cytosolic superoxide dismutase (Cu/ 7,8-dihydromethysticin contributed to CYP1A1
ZnSOD), tumor necrosis factor alpha mRNA induction. The induction of CYP1A1 indicated a
expression and cytochrome P450 (CYP) 2E1 and potential interaction between kava or kavalactones
1A2, suggesting adaptation to oxidative stress and CYP1A1-mediated chemical carcinogenesis.
and possible drug-drug interactions. Yamazaki Weiss et al. (2005) demonstrated P-glycoprotein
et al. (2008) found that a high dose (equivalent to inhibitory activity of kava-kava and its
approximately 380 mg kavalactones/kg/day; 100 kavalactones kavalactones kavain, dihydrokavain,
times of the suggested dosage for human use) of methysticin, dihydromethysticin, yangonin and
two different types of kava products for 8 days desmethoxyyangonin in P-glycoprotein-over-
significantly increased liver weights in rats. expressing cell line P388/dx and the correspond-
CYP1A2 mRNA expression was moderately ing cell line P388. The crude extract and the
increased (2.8–7.3 fold). More importantly, the kavalactones showed a moderate to potent inhibi-
high dose of kava markedly enhanced CYP1A1 tory activity with f2 (concentration needed to
mRNA expression (75–220 fold) as well as double baseline fluorescence) values of 170 μg/ml
ethoxyresorufin O-deethylase activities and and 17–90 μM, respectively. The f2 value of yan-
CYP1A1 immunoreactivities. Thus, no observed gonin could not be determined due to its higher
adverse effect levels of kavalactones would be lipophilicity. Mathews et al. (2005) found that
lower than 380 mg/kg/day. They advised that kava could cause adverse drug reactions via inhi-
considerable attention should be paid to the pos- bition of drug metabolism. The 7-day pretreat-
sibility that kava products could induce hepatic ment with kava extract only modestly induced
CYP1A1 expression in human especially in sen- hepatic cytochrome P450 activities. The human
sitive individuals. hepatic microsomal P450s most strongly inhib-
Gene expression profiling in male B6C3F1 ited by kava extract (CYP2C9, CYP2C19,
mouse livers administered kava extract by gavage CYP2D6, CYP3A4) were inhibited to the same
for 14 weeks identified the differentially degree by a “composite” kava formulation com-
expressed drug metabolizing genes in response to posed of the six major kavalactones contained in
kava treatments (Guo et al. 2010). It was found the extract. Ki values for the inhibition of CYP2C9
that the levels of significant numbers of genes and CYP2C19 activities by methysticin, dihydro-
involving drug metabolism were changed and methysticin and desmethoxyyangonin ranged
that the pathways involving xenobiotics metabo- from 5 to 10 μM. Kava extract and kavalactones
lism, Nrf2-mediated oxidative stress response, (< or =9 μM) modestly stimulated P-glycoprotein
mitochondrial functions and others were altered. ATPase activities. Two cases of patients seen on
Their results indicated that kava extract can the psychiatric emergency and consult service
significantly modulate drug metabolizing who developed severe side effects from psycho-
enzymes, potentially leading to herb-drug inter- tropic medications in the context of kava use were
actions and hepatotoxicity. Studies showed that reported by Toohey et al. (2013). In both cases,
kava extract displayed a concentration-dependent kava use may have affected the metabolism of the
effect on hepatic CYP1A1 induction (Li et al. psychotropic medications, leading to serious side
2011). Among the six major kavalactones, effects. Growing research indicated that kava
Piper methysticum 173
most likely altered concentrations of co-adminis- vidual injection. Dihydrokawain and desme-
tered psychotropics possibly by inhibiting cyto- thoxyyangonin levels remained the same as those
chrome P450 enzymes. established for their individual administration.
Methane chemical ionization (CI) gas chroma-
tography–mass spectrometry (GC–MS) was used
Pharmacokinetic Studies by Duffield et al. (1989a) to identify some of the
human urinary metabolites of the kava lactones
After the administration of kava pyrones to rats, following the ingestion of aqueous kava extract.
about half the dose (400 mg/kg, p.o.) of dihydro- All seven major, and several minor, kava lactones
kawain was found in the urine in 48 h, about two- were identified in human urine. Observed meta-
thirds of this was hydroxylated metabolites (three bolic transformations include the reduction of the
mono- and three di-hydroxylated derivatives), of 3,4-double bond and/or demethylation of the
which p-hydroxydihydrokawain was the most 4-methoxyl group of the α-pyrone ring system.
abundant (Rasmussen et al. 1979). The remain- Demethylation of the 12-methoxy substituent in
ing third consisted of metabolites formed by yangonin (or alternatively hydroxylation at C-12
scission of the 5,6-dihydro-alpha-pyrone ring of desmethoxyyangonin) was also observed. In
and included hippuric acid (9–13 % dose). contrast to the situation prevailing in the rat, no
Lower amounts of urinary metabolites were dihydroxylated metabolites of the kava lactones,
excreted when kawain was administered, but or products from ring opening of the 2-pyrone
both hydroxylated and ring-opened products ring system, were identified in human urine.
were formed. Methysticin afforded only small Abourashed and Khan (2000) used fungal models
amounts of two urinary metabolites formed by to mimic the mammalian metabolism of two major
demethylenation of the methylenedioxyphenyl kava styryl alpha-pyrones, d-kawain and d-methys-
moiety. Urinary metabolites of the alpha-pyrones, ticin. The culture broth of Cunninghamella elegans
7,8-dihydroxyangonin and yangonin were yielded 4′-hydroxykawain from d-kawain, the
formed via omicron-demethylation. No ring- same metabolite identified in rat urine. The fungus
opened products were detected. These lipophilic Torulopsis petrophilum biotransformed d-methys-
kava pyrones had extremely low solubility in ticin to 3′-hydroxy-4′-methoxykawain which was
water, which would be expected to reduce their analogous, but not identical, to a known rat metab-
absorption rates and appeared to be responsible olite of methysticin. Kavalactones appeared to be
for the variable and low extent of metabolism hydroxylated by the cytochrome P450 system and
observed. their metabolism may be enhanced by the presence
Intraperitoneal (i.p.) administration of mice of glutathione (Russmann et al. 2001; Tinsley
with 100 mg/kg of dihydrokawain, kawain, des- 1999).
methoxyyangonin and yangonin was investigated The oral pharmacokinetics of the kavalactone,
in relation to uptake by mouse brain (Keledjian kawain (100 mg/kg), were determined in rats
et al. 1988). After 5 min, dihydrokawain and with and without coadministration of kava extract
kawain attained maximum concentrations of 64.7 (256 mg/kg) to study the effect of the extract on
and 29.3 ng/mg wet brain tissue, respectively. drug disposition (Mathews et al. 2005). Kawain
Desmethoxyyangonin and yangonin had poorly was well absorbed, with >90 % of the dose elimi-
defined maxima corresponding to concentrations nated within 72 h, chiefly in urine. Compared
of 10.4 and 1.2 ng/mg wet brain tissue, respec- with kawain alone, co-administration with kava
tively, and these compounds were more slowly extract caused a tripling of kawain AUC (0–8 h)
eliminated from brain tissue. When crude kava and a doubling of Cmax. However, a 7-day pre-
resin was administered i.p. at a dose of 120 mg/ treatment with kava extract (256 mg /kg/day) had
kg, the concentrations of kawain and yangonin no effect on the pharmacokinetics of kawain
markedly increased (2 and 20 times, respectively) administered on day 8. Studies by Fu et al. (2012)
relative to the values measured from their indi- found among the six major kavalactones (kavain,
174 Piperaceae
methysticin and desmethoxyyangonin) and high-dose dogs, Kavaform was mildly toxic; pro-
their respective metabolites (p-hydroxykavain, liferation of the small cells of the thyroid epithe-
m,p-dihydroxykavain and p-hydroxy-5,6- lium and a multicentric necrosis of the
dehydrokavain), p-hydroxykavain and m,p-di- parenchyma of the liver were observed as a single
hydroxykavain were the only metabolites histological finding in one high-dose dog. Hsu
detected in the perfusate of rat liver. Kavalactone et al. (1994) tested dihydrokavain and desme-
biliary excretion was negligible. Kavalactones thoxyyangonin in mice and Wistar rats on oral
were found to be potentially bioavailable as they application. An application of 30, 100 or 300 mg/
all readily crossed the Caco-2 cell monolayers kg twice a day for 2 weeks did not yield any clini-
with apparent permeabilities (Papp) increasing cally relevant changes in haematological and his-
from 42 × 10−6 cm/s and most exhibiting more tological parameters. Rats were administered
than 70 % crossing within 90 min (Matthias et al. similar dosages over 3 months. Only with the use
2007). Not all differences in their bioavailability of desmethoxyangonin, transient changes of
can be related to kavalactone structural differ- serum glucose, cholesterol and triglycerides were
ences as it appeared that bioavailability may also found, however, without a clear dosage–effect
be affected by co-extracted compounds. For relationship. Overall, the experiment did not
example, the Papp for kawain from ethanol extracts yield hints on toxic effects of kavalactones.
was higher than the values obtained for the same Feeding Wistar rats of both sexes with 7.3 or
compound from water extracts or for the kavalac- 73 mg/kg body weight of ethanolic kava extract
tone alone. The pilot study by Tarbah et al. (2013) for 3 and 6 months elicited no signs of toxicity in
found that kavalactones (kavain; 7,8-dihydrokavain, relation to changes in body weight, hematologi-
methysticin; 7,8-dihydromethysticin; 5,6-dehydro- cal and liver parameters, and macroscopical and
kavain (=desmethyoxyyangonin); and yangonin) microscopical histological changes in the major
accumulated in the keratin matrix of hair and could organs (Sorrentino et al. 2006).
provide an easily applicable system for assessing None of the six major kavalactones was found
chronic consumption of kava. to be positive in the experimental concentration
ranges tested by the umu test (a sensitive test for
point mutations) (Jhoo et al. 2007). However,
Toxicological Studies among the different solvent fractions, the
n-butanol fraction of kava leaves was positive for
Oral administration of a test mixture of 40 % mutagenicity. Further investigations using
kavain, 40 % dihydrokavain, and 20 % yangonin bioassay-directed isolation and analysis indicated
at doses of 100 or 500 mg/kg on days 6–15 of that 2 C-glycoside flavonoid compounds identi-
gestation.was found not teratogenic or embryo- fied as 2″-O-rhamnosylvitexin and schaftoside
toxic in Wistar rats (Hapke et al. 1971). The mix- accounted for the positive mutagenic results.
ture was also negative for teratogenic activity in Whittaker et al. (2008) evaluated the toxicity and
New Zealand strain rabbits when administered mutagenicity of two commercial samples of
orally at doses of 20 or 200 mg/kg on days 6–18 kava, Kaviar and KavaPure, and the six pure kav-
after mating. However, there was a significant alactones, kawain, dihydrokawain, methysticin,
dose-related reduction of foetal weight in treated dihydromethysticin, yangonin and desmethoxyy-
rabbits. In subchronic toxicological studies of angoni including both d-kawain and dl-kawain,
Kavaform®, a German geriatric preparation con- in L5178Y mouse lymphoma cells. They found
taining 50 mg of d,1-kavain and 200 mg of mag- neither the kava samples nor the kavalactones
nesium orotatein in Wistar rats and mongrel induced a mutagenic response in the L5178Y
dogs, serum glutamate pyruvate transaminase mouse lymphoma mutation assay with the addi-
(SGPT) levels were significantly increased in tion of human liver S9 activation.
high-dose rats; however, liver cell damage was Under the conditions of the 2-year toxicology
not confirmed by histological examination. In and carcinogenesis studies of kava kava extract
Piper methysticum 175
(CAS No. 9000-38-8) in F344/N rats and B6C3F1 severe hepatotoxic reactions to kava. Incubation
mice (gavage studies) by US Department of of kava lactones (methysticin, yangonin and des-
Health and Human Services, National Toxicology methoxyyangonin) and an ethanolic extract of
Program, there was equivocal evidence of carci- dried kava root in culture with MCL-5 cells, a
nogenic activity of kava kava extract in male human lymphoblastoid cell line stably trans-
F344/N rats based on marginal increases in the fected with five human P450’s (CYP 1A1, 1A2,
incidences of testicular interstitial cell adenoma 2A6, 2E1 and 3A4) and human epoxide hydro-
(NTP 2012). There was no evidence of carcino- lase and a control cell line (cH2) derived from the
genic activity of kava kava extract in female same parental line as MCL-5, but transfected
F344/N rats administered 0.1, 0.3 or 1.0 g/kg. with two empty vectors, elicited varying degrees
There was clear evidence of carcinogenic activity of metabolic toxicity (IC50 values ranged from
of kava kava extract in male B6C3F1 mice based 50 to > 100 μM) to both MCL-5 and cH2 cell
on increased incidences of hepatoblastoma. lines; however, both cell lines were equally sensi-
There was some evidence of carcinogenic activ- tive to the test compounds (Zou et al. 2004a). The
ity of kava kava extract in female B6C3F1 mice results suggested that the parent compound for
based on increased incidences of hepatocellular each of the four test compounds was primarily
adenoma or carcinoma (combined). In gentic responsible for the observed cell toxicity and that
toxicology, kava kava extract was negative in CYP 1A1, 1A2, 2A6, 2E1 and 3A4 or epoxide
Salmonella typhimurium strains TA97, TA98, hydroxylase did not appear to be involved. It was
TA100 and TA1535 with and without S9 activa- concluded that in-vitro kava did not appear to be
tion and also negative in Escherichia coli WP2 activated to toxic metabolites by enzymes known
uvrA/pKM101 with and without S9. In-vivo, no to be important in metabolic toxicity. Studies by
increases in the frequencies of micronucleated Whitton et al. (2003) found that the extraction
erythrocytes were observed in peripheral blood process (aqueous vs. acetone in the two types of
of male or female B6C3F1 mice administered preparations) was responsible for the difference
kava kava extract by gavage for 3 months. in toxicity as extraction of glutathione in addition
to the kava lactones was important to provide
protection against hepatotoxicity. The Michael
Kava Hepatotoxicity reaction between glutathione and kava lactones,
resulting in opening of the lactone ring, reduced
In-Vitro Studies the side effects of the kava kava extracts. This
Exposure of human hepatoma cells, HepG2, to protective activity was demonstrated using
100 μM pipermethystine caused 90 % loss in cell Acanthamoebae castellanii in which 100 % cell
viability within 24 h, while 50 μM caused 65 % death occurred with 100 mg/ml kava lactones
cell death (Nerurkar et al. 2004). Similar concen- alone, and 40 % cell death with a mixture of
trations of kavalactones (7,8-dihydromethysticin 100 mg/ml glutathione and 100 mg/ml kava
and desmethoxyyangonin) did not affect cell via- lactones.
bility for up to 8 days of treatment. Mechanistic Jhoo et al. (2006) found that organic solvent
studies indicated that, in contrast to kavalactones, fractions displayed a much stronger cytotoxicity
pipermethystine significantly decreased cellular than water fractions for kava root, leaf and
ATP levels and mitochondrial membrane poten- stem peelings as evaluated in HepG2 cells
tial and induced apoptosis as measured by the based on the 3-(4,5-dimethylthiazol-2-yl)-2,5-
release of caspase-3 after 24 h of treatment. diphenyltetrazolium bromide assay and lactate
These observations suggested that pipermethys- dehydrogenase and aspartate aminotransferase
tine, rather than kavalactones, was capable of enzyme leakage assays. The hexane fraction of
causing cell death, probably in part by disrupting the root exhibited stronger cytotoxic effects than
mitochondrial function. Thus, it was concluded fractions of root extracted with other solvents
that pipermethystine may contribute to rare but (ethyl acetate, n-butanol and water) or extracts
176 Piperaceae
from the other parts of kava. Further investiga- root, to be a potent hepatocellular toxin, inducing
tions using bioassay-directed isolation and analy- cell death in HepG2 (LD50 = 15.3 μM) and l-02
sis of the hexane fraction indicated that the (LD50 = 32 μM) cells. Hepatocellular toxicity of
compound responsible for the cytotoxicity was FKB was mediated by induction of oxidative
flavokavain B. stress, depletion of reduced glutathione (GSH),
The pyridone alkaloid pipermethystine had inhibition of IKK activity leading to NF-κB tran-
been considered to be responsible for alleged scriptional blockade and constitutive TNF-α-
hepatoxicity of Kava products (Lechtenberg et al. independent activation of mitogen-activated
2008). Investigation of a series of retain samples protein kinase (MAPK) signaling pathways,
of finished products from the German market and namely, ERK, p38 and JNK. They further dem-
self-produced extracts from root and stem mate- onstrated by non-invasive bioluminescence imag-
rial of Piper methysticum clearly showed that ing that oral consumption of FKB leads to the
pipermethystine was absent from all root and inhibition of hepatic NF-κB transcriptional activ-
retain samples and extracts, with a limit of quan- ity in-vivo and severe liver damage.
tification of 45 ppm. Thus they stated that piper- Hepatotoxicity could occur as an acute, severe
methystine should not be the responsible form or a chronic, mild form in relation to kava
constituent in hepatoxic case reports with ethano- ingestion (Rowe et al. 2011). Inflammation
lic kava extracts produced in Germany. appeared to be involved in both forms and may
Studies by Lüde et al. (2008) found kava result from the activation of liver macrophages
extracts were toxic to mitochondria, leading to (Kupffer cells), either directly or via kava metab-
inhibition of the respiratory chain, increase in olites. Pharmacogenomics may influence the
ROS production, decrease in the mitochondrial severity of this inflammatory response. According
membrane potential and eventually to apoptosis to Clouatre (2004), three possible mechanisms
of exposed HepG2cells. The methanolic and an for kavalactone hepatotoxicity were known: inhi-
acetonic kava root and a methanolic leaf extracts bition of cytochrome P450, reduction in liver glu-
showed cytotoxicity starting at a concentration of tathione content and, more remotely, inhibition
50 μg/ml (lactate dehydrogenase leakage) or of cyclooxygenase enzyme activity. The patho-
1 μg/ml (MTT test). The mitochondrial mem- physiology of kava hepatotoxicity remains
brane potential was decreased (root extracts start- inclonclusive despite the existence of circum-
ing at 50 μg/ml) and the respiratory chain stantial evidence for the roles of toxic metabo-
inhibited and uncoupled (root extracts) or only lites, inhibition of cyclooxygenase (COX)
uncoupled (leaf extract) at 150 μg/ml, and mito- enzymes and depletion of liver glutathione
chondrial beta-oxidation was inhibited by all (Zhang et al. 2011). Experimental and clinical
extracts starting at 100 μg/ml. Induction of apop- cases of hepatotoxicity showed evidence of
tosis was demonstrated by all extracts at a con- hepatitis.
centration of 150 μg/ml. They stated that in
predisposed patients, mitochondrial toxicity of Animal Studies
kava extract may explain hepatic adverse reac- Studies by Singh and Devkota (2003) found that
tions of this drug. Kava was found to potentiate aqueous kava extract did not affect liver function
acetaminophen-induced hepatocyte cytotoxicity in rats. The data showed that none of the enzymes
(Yang and Salminen 2011). The findings indi- such as alanine aminotransferase, aspartate ami-
cated that kava potentiated acetaminophen- notransferase, alkaline phosphatase,lactate dehy-
induced cytotoxicity of rat primary hepatocytes drogenase andmalondialdehyde were elevated; in
by increasing the magnitude of glutathione deple- fact in some cases they were significantly
tion, resulting in oxidative stress and mitochon- reduced, suggesting the lack of a toxic effect by
drial dysfunction, ultimately leading to cell death. kava on the liver. Separate studies by DiSilvestro
Studies by Zhou et al. (2010) demonstrated et al. (2007) found 3 months of kava feeding in
that flavokawain B (FKB), a chalcone from kava rats at three different doses (31.25, 62.5 and
Piper methysticum 177
133 mg/kg diet) produced no liver injury based study of human cancers. Zhang et al. (2013)
on serum markers of liver damage (sorbitol dehy- found that liver macrophage depletion amelio-
drogenase activities, bile acid concentrations and rated kavalactone damage in isolated perfused rat
beta-glucuronidase activities) and serum lipid liver.
peroxide readings. For some measurements and In prechronic studies, orally administered
some kava doses, the injury marker readings kava at 0.125–2 g/kg body weight revealed dose-
were below control values. Moreover, for these related increases in liver weights and incidences
same parameters, kava feeding did not enhance of hepatocellular hypertrophy in F344/N rats and
the effects of the hepatotoxin galactosamine B6C3F1 mice (Behl et al. 2011). In the chronic
(500 mg/kg ip); some kava doses even showed studies, there were dose-related increases in the
modest protection against liver injury. Liver his- incidences of hepatocellular hypertrophy in rats
tology analysis showed no signs of kava causing and mice administered kava kava for up to 1 g/kg
or enhancing liver injury. Their study did not sup- body weight. This was accompanied by signifi-
port the concept that kava produces or aggravates cant increase in incidences of centrilobular fatty
liver injury. Studies by Narayanapillai et al. change. Male mice showed a significant dose-
(2014b) found that kava alone revealed no related increase in the incidence of hepatoblasto-
adverse hapatoxic effects for long-term usage mas. In female mice, there was a significant
even at a dose of 500 mg/kg bodyweight in increase in the combined incidence of hepatocel-
C57BL/6 mice. In contrast, a 3-day kava pre- lular adenoma and carcinoma in the low- and
treatment potentiated acetaminophen P-induced mid-dose groups but not in the high-dose group.
hepatotoxicity, resulted in an increased serum These findings were accompanied by several
ALT and AST and increased severity of liver nonneoplastic hepatic lesions.
lesions. It was found that flavokawains A and B
in kava, not dihydromethysticin, potentiated Clinical Studies/Reports
acetaminophen-induced hepatotoxicity. A case of a woman with acute necrotizing hepati-
Dietary feeding of flavokawain A (FKA), a tis after taking herbal remedies as alternative
major kava chalcone, did not affect food con- medication, containing kava, was reported in
sumption and body weight of male FVB/N mice Germany in 1998 (Strahl et al. 1998). Viral, auto-
(Li et al. 2014). Histopathological examination immune and metabolic causes of the hepatitis
of liver, kidney, colon, lung, heart, spleen and were excluded. Campo et al. (2002) reported a
thymus revealed no signs of FKA-induced toxic- case of a girl aged 14 years who was hospitalised
ity. Biochemical serum analysis and histological with kava-induced fulminant hepatic failure in
examination confirmed normal organ function in December 2000, after taking two kava products
FKA-treated mice. The cytotoxicity profile from August to December. Initial liver biopsy
showed FKA had minimal side effects on bone revealed active fulminant hepatitis with extensive
marrow and small intestinal epithelial cells com- centrilobular necrosis, approximately 25 % hepa-
pared with adriamycin. In addition, oral feeding tocellular viability, and mixed inflammatory infil-
of FKA increased activities of both glutathione trates consisting of lymphocytes, histiocytes,
S-transferase and quinone reductase in the liver, scattered eosinophils and occasional neutrophils.
lung, prostate and bladder tissues of mice. In No viral cytopathic changes were identified, and
comparison, dietary feeding of 0.6 % commercial immunohistochemical stains for hepatitis B sur-
kava root extract increased liver/body weight face and core antigens were negative. Humberston
ratio and decreased spleen, thymus and testis/ et al. (2003) described a case of a previously
body weight ratios, as well as induced nodular healthy 14-year-old female was admitted to the
proliferation in liver tissues. Therefore, dietary hospital with hepatic failure. Initial therapy,
feeding FKA showed no adverse effects on major including plasmapheresis, was unsuccessful and
organ function and homeostasis in mice, suggest- she deteriorated. She ultimately required a liver
ing the potential of FKA for chemoprevention transplant and now remains well. The liver biopsy
178 Piperaceae
showed hepatocellular necrosis consistent with sionflower (Passiflora incarnata). The patient
chemical hepatitis. A work-up for alternative died after a report by the Therapeutic Goods
causes of liver failure was negative. The patient Administration (TGA) warning of the potential
gave a history of taking a kava kava – containing for hepatotoxicity associated with the use of
product for 4 months. kava-containing products. In a study of 31 healthy
Russmann et al. (2003) reported that tradi- adult Tongan kava drinkers versus a control group
tional aqueous kava extracts were the most prob- of 31 healthy adult Tongan non-kava drinkers in
able cause of hepatitis in two patients presenting Oahu Hawaii, it was found that chronic heavy
with markedly elevated transaminases and hyper- kava beverage consumption was associated with
bilirubinaemia. A consequent survey of 27 heavy the elevation of γ-glutamyl transpeptidase in
kava drinkers in New Caledonia showed elevated 65 % of the kava drinkers versus 26 % in the con-
gamma glutamyl transferase in 23/27 and mini- trols (Brown et al. 2007). Alkaline phosphatase
mally elevated transaminases in 8/27. They con- was elevated in 23 % of kava drinkers versus 3 %
cluded that not only commercially available but in the controls.
also traditionally prepared kava extracts may In an otherwise healthy 48-year-old female
rarely cause liver injury. The increased activity of patient, acute hepatitis with transaminase increase
gamma glutamyl transferase in heavy kava con- (glutamic oxaloacetic transaminase (GOT) up to
sumers in the presence of normal or minimally 613 U/l, glutamic pyruvic transminase (GPT) up
elevated transaminases was probably not a sign to 752 U/l), inconspicuous hepatitis serology
of liver injury, but rather reflected an induction of findings, negative autoantibody status and nega-
CYP450 enzymes. In a cross-sectional study with tive virus serology was observed after a 10-week
98 indigenous Australian participants, 36 of long intake of kava kava (1–3 × 200 mg/day) and
whom had never used kava, it was found that St John’s Wort (1 × 425 mg/day) (Musch et al.
more recent kava use was independently associ- 2006). Biopsy of the liver showed lobular and
ated with higher levels of liver enzymes gamma- portal necroinflammatory activity without indica-
glutamyl transferase (GGT) and alkaline tion of cirrhosis. Discontinuation of the existing
phosphatase (ALP), but not with alanine amino- medication and simultaneous onset of immuno-
transferase or bilirubin, which were not elevated suppressive combination therapy of cortisone,
(Clough et al. 2003a). In those who were not azathioprine and ursodeoxycholic acid resulted
heavy alcohol users, only those who used kava in normalisation of the liver parameters within a
within the previous 24 h showed GGT levels period of 2 months. It appeared that simultaneous
higher than nonusers, whereas higher ALP levels intake of St John’s Wort possibly potentiated the
occurred only in those who last used kava 1–2 toxicity of kavapyrones. On the other hand, an
weeks and 24 h previously. It was concluded that immune-mediated mechanism, induced by kava
liver function changes in users of aqueous kava kava, could not be completely excluded. A com-
extracts at these moderate levels of consumption parative study of two structured quantitative ana-
appeared to be reversible and may begin to return lytical methods: the system of Maria and
to baseline after 1–2 weeks abstinence from kava. Victorino (MV) and that of the Council for
No evidence for irreversible liver damage had International Organizations of Medical Sciences
been found. In another paper, Clough et al. (CIOMS) for causality assessment of kava hepa-
(2003b) conducted a cross-sectional study of 101 totoxicity was carried out on 26 patients with
aboriginal adults in east Arnhem Land in the assumed hepatotoxicity by the herb (Teschke
Northern Territory. They found that kava use was et al. 2010). Grades of causality for suspected
associated with dermopathy, liver function abnor- hepatotoxicity by kava were much lower when
malities and decreased lymphocytes. Gow et al. evaluated by structured quantitative causality
(2003) described a case of acute liver failure and assessment scales than by regulatory ad hoc
death associated with the use of a preparation judgements. The quantitative CIOMS scale was
containing the “natural” anxiolytic kava and pas- found to be the preferable tool for causality
Piper methysticum 179
the formation of reactive metabolites and cova- tion to kava hepatotoxicity, they (Teschke et al.
lent binding to skin proteins leading to immune- 2011d) proposed a six-point plan for new kava
mediated responses. Reactive metabolites of kava standardization: (1) use of a noble kava cultivar
lactones had been identified in-vitro as glutathi- such as Borogu, at least 5 years old at time of
one (GSH) conjugates and in-vivo as mercaptur- harvest; (2) use of peeled and dried rhizomes and
ates excreted in urine. Unfortunately, only a few roots; (3) aqueous extraction; (4) dosage recom-
studies had investigated the toxicity of the minor mendation of ≤ 250 mg kavalactones per day (for
constituents present in kava extract, such as medicinal use); (5) systematic rigorous future
pipermethystine and the flavokavains, where research; and (6) a Pan Pacific quality control
some have been shown to display higher in-vitro system enforced by strict policing. Teschke and
cytotoxicity than the lactones. They reiterated Lebot (2011) recommended for the establishment
that to date, there remains no indisputable reason of Pan-Pacific kava quality legislation as an
for the increased prevalence of kava-induced important part of the proposed Kava Quality
hepatotoxicity in Western countries. Teschke Standardization Code. They riterated that a
et al. (2012) in a review of cases analyzed by the sophisticated approach to establish kava quality
World Health Organization and published case standardizations was needed for safe human use
reports found that traditional aqueous extracts of kava as relaxing traditional beverages, anxio-
used in New Caledonia, Australia, the USA and lytic drugs and recreational dietary supplements.
Germany may also be hepatotoxic and may not Teschke et al. (2013) reported that kava hepato-
be restricted to use of Western acetonic and etha- toxicity was presently not reproducible experi-
nolic extracts. They stated that the primary cause mentally in preclinical models, as demonstrated
of toxicity may reside in the time before the prep- by studies showing whole kava extracts were not
aration of the various kava extracts, possibly hepatotoxic. This led them to propose the
attributed to poor quality of the raw material “working hypothesis” that contaminant hepato-
caused by mould hepatotoxins. They advised for toxins including moulds might have caused rare
rigorous testing of kava raw material, in addition kava hepatotoxicity in humans and that further
to Pan-Pacific kava manufacturing quality stan- studies were warranted to proof or disproof their
dards. According to Teschke et al. (2011a;b) thus working hypothesis. Currently, there is a paucity
far no clear evidence existed for the for a caus- of evidence supporting the recent speculation
ative hepatotoxic role of kavalactones and non- that adulterants or impurities such as the mould
kavalactone constituents, such as pipermethystine hepatotoxin aflatoxin could more likely cause
and flavokavain B identified from kava. They kava hepatotoxicity rather than kava components,
asserted that studies should focus on the identifi- such as kavalactones, pipermethystine and flavo-
cation of further potential hepatotoxic constitu- kavain B, that had demonstrated hepatotoxicity
ents, considering in particular possible kava (Rowe and Ramzan 2012). Although there was
adulterants and impurities with special reference limited similarity between acute kava hepatotox-
to ochratoxin A and aflatoxins (AFs) producing icity and acute aflatoxicosis, and background lev-
Aspergillus varieties. Also due to to high temper- els of aflatoxin had been detected in kava samples,
ature and humidity in the South Pacific area, kava there is an urgent need for epidemiological inves-
raw material might have been contaminated by tigations to uncover direct evidence implicating
mould hepatotoxins such as aflatoxins after har- mould hepatotoxins.
vest and during storage (Teschke et al. 2011b).
The analysis conducted by Teschke et al. (2011c)
showed poor quality of kava raw material as a Other Adverse Health/
cause for its liver toxicity and suggested preven- Dermatological Issues
tative measures by going back to the traditional
use of kava for the sake of the patients and the In a hospital-based health survey of 150 people
South Pacific economy. Consequently as a solu- on kava drinking in Vanuatu, Grace (2003) found
Piper methysticum 181
that 59 % of men and 15 % of women drank kava review of key evidence, their findings were as
(approximately 4:1). 51 % of all men drank kava follows: causality indicated: scaly skin rash,
at least weekly, compared to 11 % of women. For weight loss, raised gamma glutamyl transpepti-
any given kava-drinking episode, men drank dase liver enzyme levels, nausea, loss of appetite
more than women, (4.3 vs 3.3 shells). There was or indigestion; association indicated but causality
no significant difference in age between drinkers unclear: red sore eyes, impotence or loss of sex-
and non-drinkers or in usage rates between ual drive, self-reported poor health, raised cho-
patient groups or staff. The study revealed that lesterol, and loss of time and money, low
the consumption of fresh kava on a regular basis motivation and “slow/lazy” days following use,
was very common in Vanuatu. Studies by Cairney reduced alcohol consumption and related vio-
et al. (2003) on a group of current, ex and non- lence; association hypothesised: fits or seizures,
kava users among an indigenous population in melioidosis, ischaemic heart disease, protective
northern Australia found no impairment in cogni- effects for cancer; no association indicated: cog-
tive or saccade function in individuals who were nitive performance; no association suggested:
currently heavy kava users (and had been for up cognitive impairment, liver toxicity or permanent
to 18 years), nor was there any impairment in liver damage and other pneumonia; no associa-
individuals who had been heavy kava users in the tion hypothesised: hallucinations. The study of
past but had abstained for longer than 6 months. Vignier et al. (2011) found that kava drinking
Current and ex-kava users showed a higher rate was associated with suicidal behaviour among
of kava dermopathy, lower body mass index, low- young Kanaks using kava in New Caledonia but
ered blood lymphocytes and, in addition, current information on the effects of kava on mental
kava users showed elevated liver enzymes. They health in young people was lacking and require
found no evidence of brain dysfunction in heavy further investigations.
and long-term kava users. In 2000 dermopathy Chronic consumption (6 months) of large
characteristic of heavy use, abnormally low body quantities of an infusion of kava rhizome (5–6
mass index (BMI), low blood lymphocytes and cups daily) had been reported to cause anorexia,
abnormally high gamma-glutamyl transferase diarrhoea and visual disturbances (Siegel 1976).
(GGT) occurred more frequently with increased In a healthy volunteer, disturbances of visual
kava use in Arnhem Land Aboriginal communi- accommodation, such as enlargement of the
ties in the Northern Territory (Clough 2003). pupils and disturbances in oculomotor equilib-
These acute effects emerged at average consump- rium, were reported following the ingestion of
tion levels of 310–440 g/week of kava powder. large doses of kava (Garner and Klinger 1985).
The proportion of men drinking kava reached No changes were recorded in visual or stereoacu-
70 % and women 62 % from mid-1990, with ity or in ocular refractive error. Mathews et al.
20 % of the population spending unprecedented (1988) conducted a pilot health survey on kava
amounts of time (14 + hours/week) in activities usage involving 20 very heavy users of kava
where kava was consumed. (mean consumption 440 g/week), 15 were heavy
Kava being used for a wide spectrum of thera- users (310 g/week) and 4 were occasional users
peutic properties, including sedative, anxiolytic, (100 g/week). They found that in addition to
analgesic and neuroprotective effects, was one of causing acute intoxication, sedation and relax-
eight herbal supplements identified that could ation, a rash and weight loss in long-term users,
pose the greatest potential risks in surgical kava may also cause liver and renal dysfunction,
patients in the United States (Sullivan et al. hematological abnormalities and possibly pul-
2009). An Australia-wide ban on commercial monary hypertension. The conclusion that kava
importation of kava has been in place since mid- affected the liver was based on the markedly ele-
2007, but published literature on the impact of vated plasma levels of γ-glutamyl transferase
the ban had been lacking to date on the impact of observed in kava users. Heavy consumption of
the ban (Rychetnik and Madronio 2011). In a kava has been reported to lead to adverse health
182 Piperaceae
effects in some Australian Aborigines including 70-year-old man and a 52-year-old woman, with
the occurrence of pellagroid dermopathy (Cawte sebotropic drug eruption in sebaceous gland
1986, 1988). Bilia et al. (2002) reviewed nine areas induced by 3 weeks of systemic kava kava
clinical trials and found that three of these trials anti-depressant therapy. Huynh et al. (2014)
reported no adverse events, while the other six reported a 55-year-old man presented with an
studies reported gastrointestinal symptoms, tired- eruption in a sebotropic distribution after con-
ness, restlessness, tremor and headache. The suming kava kava for 3 weeks, which resolved
number of patients reporting the complaints was after discontinuation of the supplement. Shimoda
similar in the placebo groups. Aside from liver et al. (2012) found that mast cells exposed to
abnormalities or toxicity, adverse effects attrib- standardised kava aqueous extracts displayed
uted to kava extracts included gastrointestinal robust intracellular free calcium responses and
complaints, restlessness, mydriasis, allergic skin concomitant release of proinflammatory media-
reactions, dermatomyositis (Ernst et al. 2001), tors. In contrast, mast cells were refractory to
visual accommodation disorders, pupil dilation single or combinatorial stimulation with kavalac-
and disorders of oculomotor equilibrium (Singh tones, including methysticin, dihydromethysticin
and Blumenthal 1997). Toxic doses (several and kavain. The study suggested that mast cell
times the therapeutic dose of approximately activation may be a mechanistic component of
70 mg of kavalactones three times daily) could kava-related skin inflammations.
cause progressive ataxia, muscle weakness and Studies by Xia et al. (2012) suggested that
ascending paralysis (Spillane et al. 1997). kava was photocytotoxic and photogenotoxic,
Heavy kava drinkers were reported to acquire both mediated by free radicals generated during
a reversible peculiar, scaly, ichthyosiform erup- photoirradiation. They found that UVA irradia-
tion on the skin and kava dermopathy (Norton tion of kava in the presence of a lipid, methyl
and Ruze 1994). Traditional kava use at frequent linoleate, generated lipid peroxidation which was
high doses had been found to cause a reversible mediated by singlet oxygen generated during
dermatological condition known as ichthyosi- photoirradiation. Of the six major kavalactones
form ichthyosiform kava dermopathy (Süss and (yangonin; 7,8-dihydrokawain; kawain,
Lehmann 1996; Dentali 1997), known as kani- 7,8-dihydromethysticin; methysticin; and
kani in Fijian (Gounder 2006). Kava dermopathy 5,6-dehydrokawain), only 5,6-dehydrokawain
was believed to be related to interference with and yangonin-induced a low level of lipid peroxi-
cholesterol metabolism (Ruze 1990; Norton and dation. UVA irradiation of kava in human HaCaT
Ruze 1994; Dentali 1997). Ruze (1990) found skin keratinocytes induced cytotoxicity which
that the pellagroid dermopathy was not related to was mediated by oxidative stress, led to DNA
niacin deficiency and was more characteristic of strand cleavage, and produced 8-hydroxy-2′-
an acquired ichthyosis. In Fiji in 2012, over 1000 deoxyguanosine (8-OHdG) adduct. Study by the
kava participants underwent full skin examina- electron spin resonance (ESR) method revealed
tion, and kava dermopathy was a common cuta- that UVA irradiation of kava produced singlet
neous finding (Hannam et al. 2014). The clinical oxygen and carbon-centered radicals.
manifestations of kava dermopathy shared simi- Schelosky et al. (1995) reported four cases of
larities with the spectrum of autosomal recessive patients who developed clinical signs suggestive
congenital ichthyoses, predominantly lamellar of central dopamine antagonisms after exposure
ichthyosis. The pathogenesis of Kava dermopa- to various kava preparations for anxiety. These
thy may be associated with a functional defect in case histories suggested that the sedative effects
one or more cytochrome P450 enzymes impli- of kava might result from dopamine antagonistic
cated in epidermal integrity, thus mimicking the properties of kava extracts. They highlighted the
genetic defect as seen in lamellar ichthyosis type pyramidal side effects of kava preparations and
3. Jappe et al. (1998) described two patients, a cautioned their use particularly in elderly patients.
Piper methysticum 183
Meseguer et al. (2002) reported a 45-year-old In Tubuai, a maceration of young kava shoot is
female with severe parkinsonism induced by taken orally for inflammation of the genito-
kava kava. The patient, who had a family history urinary system (Aitken 1930) In the Pacific
of essential tremor, developed severe and persis- Islands, kava is taken as a drink for gonorrhoea
tent parkinsonism after days of treatment with and chronic cystitis (Steinmetz 1960). In Papua
kava extract for anxiety. The symptoms improved New Guinea, kava is used as an anaesthetic and
with anticholinergics. They asserted that kava stimulant for lactation (Steinmetz 1960); the bark
derivatives could produce severe parkinsonism in of scraped and masticated root is used to relive
individuals with genetic susceptibility. sore throats, juice from leaves is used to treat cuts
A case of rhabdomyolysis temporally related and drunk as tonic (Holdsworth 1977). In certain
to the ingestion of a large amount of kava was tribes of Papua New Guinea, women drink fresh
reported by Bodkin et al. (2012) in addition to an masticated kava as an anaesthetic, when they are
earlier case of rhabdomyolysis related to the being tattooed (Steinmetz 1960). They also drink
ingestion of kava. Their patient developed peak large quantities when they are pregnant, espe-
creatine phosphokinase levels in excess of 30 000 cially just before delivery, to stimulate milk pro-
U/L but had no significant renal damage. In May duction. In Irian Jaya, the internal skin of the
2013, a case of hepatitis A infection was reported kava plant is used for toothache (Aufenanger and
to a Western Australian regional public health Hötker 1940). In Tahiti, according to Sterly
unit, with infection acquired in Fiji (Parker et al. (1970), the masticated root is considered a rem-
2014). Following this, two further cases were edy for gonorrhoea. In American Samoa, kava
linked to the index case by kava drinking and one roots together with fruit of polo fe’u (chilli,
further case was a household contact of a second- Capsicum annuum), leaves of fisoa (soap bush,
ary case. This outbreak highlighted that the prep- Colubrina asiatica) and inner bark of moli’aina
aration of kava drink and/or the use of a common (orange, Citrus sinensis) are taken internally for
drinking vessel could be a vehicle for the trans- gonorrhoea; kava leaves are taken internally for
mission of hepatitis A virus. mumu tuala uli (swollen head, sore eyes, cold
sweat, dizziness, numbness of legs), and the inner
bark with juice of fasa (screw-pine, Pandanus
Traditional Medicinal Uses tectorius) is taken internally for tulita fasia (uri-
nary tract infection) Harrington and Scotese
Kava has been used in social and ceremonial life (2001).
in the Pacific islands from ancient times for the In Hawaii, awa root is used to treat difficulties
soporific and narcotic effects (Bilia et al. 2002). in urination (Handy et al. 1934; Titcomb 1948)
In Polynesia, about thirty syndromes were treated and irritation of the genito-urinary system (Handy
with kava-based preparations (Zepernick 1972). et al. 1934); masticated awa and drinking awa are
The traditional medicinal uses of kava include employed for feminine puberty syndromes and
treatment of gonorrhoea, syphilis, and cystitis; weakness, painful migraine, headache, macera-
induction of muscle relation and sleep (Ernst tion of awa for vaginal prolapus, maceration of
et al. 2001); and treatment of boils, asthma, head- masticated awa diluted, boiled and taken orally
ache and urinary infections (Hirsch 2000). In the for general weakness (Titcomb 1948); awa-based
Pacific Oceania, kava is held in high esteem pri- medication is used for menstrual problems and
marily as a ritual offering or a ritualised form of dysmenorrhoea, and masticated awa is drank to
payment (Lebot and Cabalion 1988). To treat prevent risk of infection (Handy et al. 1934);
injury caused by fish spines, dry kava root is maceration of masticated awa is diluted, boiled
burnt and the injury exposed to the smoke and taken orally for chills and sleeping problems
generated. (Titcomb 1948: Handy et al. 1934; Handy 1940;
184 Piperaceae
Hänsel 1968); masticated awa or as a drink is sumed by women as a fortifying drink, laxative
employed for headaches (Handy et al. 1934; and diuretic. In pregnancy, small amount taken is
Handy 1940; Titcomb 1948); maceration of rhi- said to facilitate delivery. During breast feeding,
zome is taken orally for rheumatism and against awa is taken to stimulate milk production. Kava
fat intake (Handy 1940; Titcomb 1948); awa is used medicinally also for kidney and bladder
preparation is employed for the irritation of respi- troubles and as a strong sudorific. Though like
ratory tract, a maceration of rhizome is taken all other drugs, if taken in excess it has bad
orally for gastro-intestinal upsets, masticated awa results on account of its excessive action on the
is drank for pulmonary pains, awa is used for cer- skin, and may even super-induce elephantiasis,
tain skin disease and masticated rhizome as poul- that terrible complaint, so prevalent in Fijian vil-
tice against suppuration (Titcomb 1948). Awa lages. In Fiji, kava leaves are softened in fire and
drink is administered to children to calm their applied as poultice against suppurations
nervousness (Titcomb 1948), juice from rhizome (Degener 1949). Fijian kava is used in traditional
is used in medication for tuberculosis, medica- medicine against urinary tract infections and
tion containing masticated rhizome used exter- asthma (Folmer et al. 2006). Japanese newspaper
nally for leprosy (Degener 1945) and masticated in early 1985 reported virtues of Fijian yaqona
rhizome is used in poultice for skin disease as a remedy against colds and coughing (Lebot
(Hänsel 1968). Awa leaves are placed in-situ to and Cabalion 1988)
provoke abortion (Handy et al. 1934), and fumi- Ingestion of kava is also reported to aid in first
gation with leaves is used for chill and general stage of diarrhoea (Thomson 1908; Sterly 1970).
disease treatment (Handy 1940). In Motalava, Vanuatu, kava roots are used in
In Hawaii, Awa is taken as a therapy for sooth- drink to cure constipation and used for conjuncti-
ing the nerves, relaxing fatigue-stiff muscles, vitis (Vienne 1981). Eyes are washed with juice
general debility and inducing sleep and a treat- squeezed from kava leaves.
ment for excessive fat for restoring the body to
normal fitness (Titcomb 1948). Awa treatment
was also said to overcome scaly and ulcerous Other Uses
skin. Mouritz (1916) stated that an alcoholic awa
solution injected into the skin causes anaesthesia, The use of kava over the centuries is in the form
followed by paralysis of the peripheral nerves for of a traditional ethnic beverage forming an inte-
several hours. He added that both Hawaiian and gral part of social gatherings and ceremonies to
haole (foreign) doctors use it for venereal and increase kinship and amiability, especially in the
kidney diseases, and, in alcoholic solution or as Polynesian and Melanesian societies, and is
an unguent, in affections of the skin, including widely consumed on a daily basis.
leprosy. Handy et al. (1934) stated that bits of Kava leaf/root/stem extract is used as cos-
awa root chewed at frequent intervals and awa metic skin-conditioning agents at concentrations
leaves wrapped around the head were said to pro- from 0.0001 to 0.1 % (Robinson et al. 2009).
tect against contagious disease and to cure head- Data is still wanting on whether kava-derived
ache. Awa leaves, stuffed into the vagina, were ingredients would be substantially absorbed
said to induce miscarriage. Jarrett (1933) men- through the skin.
tioned that awa was extensively used in Germany Kava had been reported to have antifungal and
prior to the World War, in the manufacture of cer- herbicidal property.
tain drugs and medicines. Kava showed a strong inhibition on growth of
In Fiji, yaqona is considered a powerful dia- barnyardgrass (Echinochloa crus-galli), mono-
phoretic (Parham 1939, 1941). It is also con- choria (Monochoria vaginalis) and knotgrass
Piper methysticum 185
(Paspalum distichum), some of the most harmful Abu N, Akhtar MN, Yeap SK, Lim KL, Ho WY, Zulfadli
AJ, Omar AR, Sulaiman MR, Abdullah MP, Alitheen
paddy weeds (Xuan et al. 2003). Kava completely
NB (2014) Flavokawain A induces apoptosis in
controlled emergence of monochoria and barn- MCF-7 and MDA-MB231 and inhibits the metastatic
yardgrass at a treated dose of 0.5 and 1.0 g, process in vitro. PLoS One 9(10):e105244
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zwei neuen pyrrolidinen aus rauschpfeffer. Chem Ber
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103:2535–2540. (In German)
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Achenbach H, Wittman G (1970) Dihydrokawain-5-ol,
addition, kava significantly inhibited growth of
ein neuer alkohol aus rauschpfeffer (Piper methysti-
the five fungi: Fusarium solani, Pyricularia gri- cum Forst.). Tetrahedron Lett 11(37):3259–3262
sea, Rhizopus stolonifer, Taphrina deformans Achenbach H, Karl W, Smith S (1971) Zur gaschro-
and Thanatephorus cucumeris. The effect on R. matograpischen trennung der kawa-lactone-(+)-
5,6,7,8-tetrahydro-yangonin, ein neus kawa-lacton aus
stolonifer was the greatest and T. cucumeris and
rauschpfeffers. Chem Ber 104:2688–2693
P. grisea were the second most affected. Aqueous Achenbach H, Karl W, Regel W (1972) 11-Hydroxy-12-
extract of kava roots showed high allelopathic methoxy-dihydrokawain und 11,12-dimethoxy-
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105:2182–2187
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Aufenanger H, Hötker G (1940) Die Gende in
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brain tissue against ischemic damage in rodents. Eur
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Chrysopogon zizanioides
Chrysopogon zizanioides (L.) Roberty Cus-Cus, Cuscus Grass, Indian Couch Grass,
Khus-Khus Grass, Kuskus, Vetiver, Vetiver
Grass, Vetiveria
Synonyms
it is a strong candidate for reclamation of wet grass). It has a woody taste and a scent similar to
land soils. Dudai et al. (2006) found that vetiver khus. The syrup is used to flavour milkshakes and
could grow in a wide range of substrates, such as yoghurt drinks like lassi, but can also be used in
sandy soil, loamy sand, clay soil, crushed lime- ice creams, mixed beverages like Shirley Temples
stone, sandy clay loam and tuff/peat mixture and as a dessert topping.
under Mediterranean conditions.
Botany
Edible Plant Parts and Uses
Vetiver is a coarse, aromatic perennial grass
The roots are also used to provide an important forming large dense clumps with massive,
ingredient in curry, khasu-khasu in India. In India spongy, finely structured, fibrous root systems
it yields a fragrant oil which is used in khas syrup spreading to a depth of 4 m (Plate 1). Culms are
and khas water to flavour sherbets as well as to robust, erect, 1–2.5 m tall, and about 5 mm in
perfume soaps and cosmetics. This extract, called diameter. Leaf sheaths glabrous, lower sharply
‘khus essence’, is added to a sugar, water and cit- keeled, laterally compressed and imbricate in
ric acid syrup. In India, vetiver essence is used to fanlike clusters; leaf blades linear, pale green,
flavour a soft drink (Sharbat), syrup and ice stiff, 30–90 × 0.5–1 cm, pilose on adaxial surface
cream (Lavania 2003). Chopped, dried vetiver towards base, otherwise glabrous (Plate 2); ligule
roots can be made into potpourri by mixing them with a scarious margin. Panicle open, exserted,
with dried fragrant flower petals and spices oblong in outline (Plate 3), 15–40 cm, usually
(Chomchalow and Chapman 2003). A stimulant
drink is made from fresh rhizomes in India. In
Thailand the Thai traditional beverage ‘Nam Ya
Faek’ is made from vetiver grass (Chomchalow
and Hicks 2001; Chomchalow and Chapman
2003).
In the hilly regions of Karnataka, India, peo-
ple made use of vetiver roots to prepare refresh-
ing drinking water (Sastry 1998). Roots are used
to prepare sherbets, ice cream or soft drinks dur-
ing summer in northern India (Lavania 2003;
Pareek and Kumar 2013). Vetiver oil is used for
flavouring syrups and ice cream, and in food
preservation. Khus essence is used in cool drinks
and for reducing pungency of chewing tobacco
preparations and to provide sweet note to other
masticatories. Vetiver roots are used domestically
in cooking; it is infused in tea and also used in
baking (Balasankar et al. 2013). In certain canned
foods, e.g. asparagus and peas, fractions of veti-
ver oil are used to reinforce the natural odour and
taste (De Guzman and Oyen 1999).
Vetiver grass is used as a flavouring agent,
usually through khus syrup. Khus syrup is made
by adding khus essence to sugar, water and citric
acid syrup. Khus essence is a dark green thick
syrup made from the roots of khus grass (vetiver Plate 1 Vetiver grass large dense clump
Chrysopogon zizanioides 201
Nutritive/Medicinal Properties
Root Phytochemicals
of vetiver oil (Chiurdoglu and Delesemme 1961). the same oil. Two tricyclic sesquiterpene acids
The structure of the bicyclovetivenol was khusenic acid and isokhusenic acid were isolated
elucidated as 2-methylene-6-methyl-9-(β- from vetiver oil (Nigam and Komae 1967).
hydroxyisopropyl)-bicyclo-(0,3,5)-decane and Khusimene was isolated from Japanese vetiver
that of veticadinol as 2-methylene-8-methyI-(β- oil (Morikawa and Hirose 1967). A laevorotatory
hydroxyisopropyl)-bicyclo-5-(0,4,4)-decane. antipodal sesquiterpene epoxy alcohol, khusino-
Dihydrobicyclovetivene and dihydroveticadinene loxide, was isolated from North Indian vetiver oil
were also obtained by dehydration of the corre- (Seshadri et al. 1967). Zizanoic acid has been
sponding dihydro-alcohols. The structures of two identified as a constituent of vetiver oil from
diolefins and two olefins were also determined. A Zambia (Grove et al. 1968). Jentsch and Triebs
laevorotatory crystalline sesquiterpene alcohol (1968a, b) isolated tertiary vetivenols: tert-
laevojunenol and a dextrorotatory alcohol junenol bicyclovetivenol and tert-tricyclovetivenol. A
were isolated from North Indian vetiver oil sesquiterpenic tricyclic primary alcohol named
(Bhattacharyya et al. 1960; Shaligram et al. khusenol was isolated from Angola vetiver oil
1962). The absolute configuration of these two (Nigam et al. 1968b). α-Vetivone (isonootka-
alcohols, from the eudesman group of com- tone) (Marshall and Andersen 1967) and
pounds, has been determined on the basis of the β-vetivone (Marshall and Johnson 1970) were
synthesis of dihydrojunenol from santanolide ‘c’. identified in Haiti vetiver oil. Tricyclic sesquiter-
The dextrorotatory alcohol junenol has also been penes khusenol, tricyclovetivenol and khusimol
similarly synthesised from the ten-membered were reported from Congo and South Indian veti-
carbocyclic lactone costunolide (Shaligram et al. ver oils (Nigam et al. 1968a). Minor sesquiter-
1962). A sesquiterpene hydrocarbon, isobisabo- pene acids zizanoic acid, isovalencenic acid,
lene (Kalsi et al. 1962); antipodal sesquiterpene epizizanoic acid (Hanayama et al. 1968), khuse-
cadinenic alcohol, khusol (Khalsi et al. 1963); a nic acid and isokhusenic acid (Komae and Nigam
laevorotatory aldehyde khusilal (Kalsi et al. 1968) were identified in vetiver oil. A bicyclic
1964); antipodal sesquiterpene alcohol cadinane, primary alcohol was isolated from vetiver oil; its
eudesmane (Kalsi et al. 1979a); cadinane ter- structure elucidated it to be a member of the ere-
penoids, khusinoloxide and epi-khusinol (Kalsi mophilane series, by correlation with α-vetivone
and Talwar 1981); C14 keto-alcohol khusitoneol, (isonootkatone) (Takahashi 1968). Three aro-
a hydroxyl derivative of khusitone (Kalsi et al. matic sesquiterpene hydrocarbons α-calacorene,
1985a); and antipodal C14 terpenoid norkhusino- dehydro-curcumene and a cadalene-type hydro-
loxide (Kalsi et al. 1985b) were isolated from carbon sesquiterpene were isolated from Réunion
Indian vetiver oil. A sesquiterpene hydrocarbon, vetiver oil (Mizrahi and Nigam 1969). An
(−)γ2-cadinene was isolated from North Indian unnamed acidic constituent with a kusilane skel-
vetiver oil (Kartha et al. 1963). Khusinol, a crys- eton was found in vetiver oil (Kalsi and Bhatia
talline secondary sesquiterpene alcohol, was iso- 1969). Two sesquiterpene primary alcohols
lated from North Indian vetiver oil (Rao et al. khusimol (Umarani et al. 1969b) and isokhusi-
1963). It was found to belong to the unusual mol (Umarani et al. 1969a, b) and a C14 ketone
antipodal group of cadinenes. A sesquiterpene khusimone (Umarani et al. 1970) were isolated
zizanoic acid (Kido et al. 1967) and minor acids from Indian vetiver oil. Minor sesquiterpene
cyclocopacamphenic and epicyclocopacam- alcohols vetiselinenol, zizanol, cyclocopacam-
phenic acids (Kido et al. 1969) were found in phenol and epicyclocopacamphenol besides
vetiver oil. A laevorotatory ketone, khusitone khusimol, valerianol (kusunol), β-eudesmol and
(Trivedi et al. 1964) and γ-cadinene, α-muurolol elemol were found in vetiver oil (Homma et al.
(old name δ-cadinol) and khusimol (Trivedi et al. 1970). MacSweeney et al. (1970) reported the co-
1971) were isolated from North Indian vetiver occurrence of tricyclovetivene, tricyclovetivenol
oil. The authors reported that γ-cadinene co- and zizanoic acid with α-vetivone and β-vetivone
occurred with γ2-cadinene, another constituent of in vetiver oil. A rare nor-terpenoid hydrocarbon
Chrysopogon zizanioides 203
with khusilane framework was isolated from North Indian vetiver oil (Karkhanis et al. 1978).
North Indian vetiver oil (Kalsi 1970). The follow- Khusinol and allokhusinol were found in vetiver
ing sesquiterpenes were reported in Reunion and oil (Ganguly et al. 1978a; b). Zizanal and
Haitian vetiver oils: α-vetivone, β-vetivone, epizizanal were isolated from Javanese vetiver oil
khusinol, khusimyl acetate elemol, β-eudesmol, (Jain et al. 1982).
vetiselinenol, 2-keto-4,10-epi-eudesmane, laevo- Several rare phenols, including 4-vinylphenol,
junenol and zizanene ((+)-α-amorphene), 4-vinyl-2-methoxyphenol and trans-isoeugenol,
prezizaene, zizanol, agarospirol, α-vetisperene, were identified in oil of vetiver (Shibamoto and
β-vetisperene, β-vetivenene, vetiselinol, 2-keto- Nishimura 1982). The tricyclic ketone 2-epi-
5-hydroxyeudesmane, anhydro-β-rotunol, hine- zizanone was isolated from Indian vetiver oil
sol, nootkatone, (−)-selina-4(14),7(11)-diene and (Moosanagar variety) (Bhatwadekar et al. 1983).
(−)-δ-selinene (Andersen 1970a, b, c; Andersen An antipodal sesquiterpene diol, vetidol, was iso-
et al. 1970; Andersen and Falcone 1971). North lated from vetiver oil (Kalsi and Talwar 1987).
Indian variety vetiver oil was found to be Oxidation of the sesquiterpenic fraction of
distinctly different from that of Reunion and Bourbon vetiver essential oil (BVEO) with
Haiti and may represent a chemically distinct m-chloroperbenzoic acid afforded three new oxi-
race of Vetiveria zizanioides species (Andersen dised products, including a zizaene monoepoxide
1970a). A C14 hydrocarbon, (+)-khisitene was and two diepoxides derived from zizanene and
isolated from North Indian vetiver oil (Raj et al. valencene (Bombarda et al. 1996). Reduction
1971). The sesquiterpene alcohol epi-khusinol reaction of sesquiterpenic epoxides of BVEO led
(Kalsi et al. 1972) and isokhusinoloxide, a new to three sesquiterpenols, including two zizanol
sesquiterpene epoxy alcohol (Kalsi et al. 1979b), epimers and an epoxialcohol derived from valen-
were found in North Indian vetiver oil. Two cene, which has been synthesised in high yields
new C12-ketones, (+)-(1S, 10R)-1,10- (85–100 %).
dimethylbicyclo[4.4.0] dec-6-en-3-one and Four new sesquiterpene ethers, 7β,10β-
(+)-(6S, 10S)-6,10-dimethylbicyclo[4.4.0]dec-1- epoxy-4βH-eremophila-1,11(12)-diene; 10β,
en-3-one, were isolated from Reunion vetiver oil 11-epoxy-4βH-eremophil-1-ene; 4α,7-epoxy-
(Maurer et al. 1972). Structure and absolute con- 10βH-spirovetiva-2,11(12)-diene; and 6α,12-
figuration of the former compound were estab- epoxy-7βH,10βH-spiroax-4-ene, were isolated
lished by a four-step synthesis from from the unpolar part of the Haitian vetiver oil
(+)-isonootkatone ((+)-α-vetivone), while the (Weyerstahl et al. 1996). The new cis-eudesma-
absolute configuration of the latter was estab- 6,11-diene was isolated from vetiver oil as a main
lished by chemical correlation with constituent of the sesquiterpene hydrocarbon
(+)-α-eudesmol. Kaiser and Naegeli (1972) iden- fraction (Weyerstahl et al. 1997). Further, six
tified 10-epi-γ-eudesmol; β-bisabolol; acora- derivatives, 13-nor-cis-eudesm-6-en-11-one; cis-
4,9-diene; acora-4,7-diene; α-cedrene; eudesm-6-en-11-ol; cis-eudesm-6-en-12-al (2
α-cedrenol; α-cedrenal; α-funebrene; epimers); 13-nor-cis-eudesm-6-en-8-one; and
α-funebrenol; α-funebrenal; and α-funebrenic cis-eudesma-6,11-dien-3-ol, were identified in
acid in vetiver oil. Two sesquiterpene carboxylic the oil. The following (nor-)sesquiterpene alco-
acids, zizanoic and epizizanoic acids, were iso- hols were isolated as their methyl ethers from
lated from the essential oil of vetiver (Hanayama the polar part of commercial Haitian vetiver
et al. 1973). Minor sesquiterpenes acora-4,9-di- oil and identified as 12-nor-ziza-6(13)-en-2β-;
ene; acora-4,7-diene; α-cedrene; α-cedrenol; and -2α-ol; eudesma-4(15),7-dien-3β-; and
α-cedrenal; α-funebrene; α-funebrenol; -2β-ol; eudesma-3,5-dien-1α-ol; eremophila-
α-funebrenal; and α-funebrenic acid were found 1(10),4(15)-dien-2α-ol; eremophila-1(10),11-
in Indian vetiver oil (Paknikar et al. 1975). Minor dien-2α-ol (nootkatol); guaia-1(5),11-dien-3-ol;
sesquiterpene alcohols isovalencenol, vestise- spirovetiva-3,7(11)-dien-12-ol; preziza-7(15)-
linenol and isovestiselinenol were isolated from en-3α-ol; and helifol-1-en-14-ol (syn. khusien-
204 Poaceae
(1.2 %), α-vetivone (6.6 %) and β-vetivone tent (Wang et al. 2009). (2-Isopopenyl-decahyero-
(5.2 %). In vetiver roots, most of the volatiles in 4,7-methano-azulen-8-yl)-methanol and other
roots were sesquiterpenes; the main component terpene compounds accounted for 11.4 %.
was valencene (30.36 %) (Huang et al. 2004). Isoeugenol and vanillin were detected in the oil
Nitrogen-containing compounds (33) of vetiver for the first time.
oil comprised substituted pyridines, pyrazines Six compounds, 5,10-pentadecadiyn-1-ol,
and indolines; two were tentatively identified as α-curcumene, hydroxy junipene, (+) cycloisosa-
isomers of trimethylindoline and isocyanato- tivene, valencine and selino 3,7 (11)-diene, were
methylbenzene (Clery et al. 2005). isolated from vetiver oil (Gupta et al. 2011).
Fourteen compounds were identified in veti- Chou et al. (2012) identified 25 components of
ver root essential oil from Bangladesh (Bhuiyan vetiver oil and the major components as cedr-8-
et al. 2008): 2,6-dimethyl-10-methylene-12-- en-13-ol (12.4 %), α-amorphene (7.80 %),
oxatricyclo[7.3.1.0(1,6)]tridec-2-ene (21.34 %); β-vatirenene (5.94 %) and α-gurjunene (5.91 %).
2-(4a,8-dimethyl-1,2,3,4,4a,5,6,7-octahydro- The major components of Brazilian vetiver
naphthalen-2-yl)-prop-2-en-1-ol (12.44 %); essential oil were identified as khusimol
caryophyllene oxide (5.78 %); juniper camphor (19.57 %), E-isovalencenol (13.24 %), α-vetivone
(3.49 %); 2-methylenecholestan-3-ol (7.13 %); (5.25 %), α-cadinol (5.01 %), β-vetivone
τ-muurolol (7.54 %); tricyclo[8.6.0.0(2,9)] (4.87 %), hydroxy-valencene (4.64 %), junenol
hexadeca-3,15-diene, trans-2,9-anti-9,10- (3.58 %), 2-epi-ziza-6(13)-em-3-ol (2.79 %) and
trans-1,10- (3.34 %); androstan-17-one; 3-ethyl- mustakone (2.75 %) (Lima et al. 2012). Other
3-hydroxy-, (5α)- (4.95 %); 1H-cycloprop[e] minor constituents (<2.5 %) were vetivenic acid,
azulen-7-ol, decahydro-1,1,7-trimethyl-4- nootkatone, spirovetiva-3,7-(11)-dien-12-ol, (Z)-
methylene-,[1ar-(1aα,4aα,7β,7aβ,7bα)]- β-curcumen-12-ol, nootkatol, junicedranol,
(4.44 %); β-vatirene (4.75 %); 3a,7- 1,7-diepi-α-cedranal, cubenol, 10-epi-γ-
methano-3aH-cyclopentacyclooctene, eudesmol, β-atlantol, khusimone, viridiflorol,
1,4,5,6,7,8,9,9a-octahydro-1,1,7-trimethyl- maaliol, β-vetivenene, elemol, γ-amorphene,
,[3aR-(3aα,7α,9aβ)]- (9.59 %); β-vetispirene, γ-gurjunene, khusimene and
2(3H)-naphthalenone, 4,4a,5,6,7,8-hexahydro- prezizaene. Eighty constituents, representing
4,4a-dimethyl-6-(1-methylethenyl)- (4.03 %); 94.5–97.8 % of the oils, were identified in Vetiver
2(1H)naphthalenone, 3,5,6,7,8,8a-hexahydro- oils from Bangalore, Hyderabad, Kundapur and
4,8a-dimethyl-6-(1-methylethenyl)- (7.78 %); Mettupalayam in South India (Mallavarapu et al.
and naphthalene, 1,2,3,5,6,7,8,8a-octahydro- 2012). The oils were found to be rich in sesqui-
1,8a-dimethyl-7-(1-methylethenyl)-, [1R- terpenes and oxygenated sesquiterpenes with
(1α,7β,8aα)]- (3.42 %). The Vetiver roots from cedrane, bisabolane, eudesmane, eremophilane
Banthra near Lucknow on hydrodistillation and zizaane skeletons. The main components of
afforded 0.28 % oil on dry weight basis the four essential oils were eudesma-4,6-diene
(Chowdhury et al. 2009). The oil contained ses- (δ-selinene) + β-vetispirene (3.9–6.1 %);
quiterpene and sesquiterpene derivatives mainly β-vetivenene (0.9–9.4 %); 13-nor-trans-eudesma-
γ-cadinene, clovene, α-amorphene, 4(15),7-dien-11-one + amorph-4-en-10-ol (5.0–
aromadendrene, junipene, β-himachalene, farne- 6.4 %); trans-eudesma-4(15),7-dien-12-ol
sene, β-bisabolene, cis-caryophyllene, khusimol, (vetiselinenol) + (E)-opposita-4(15),7(11)-dien-
epiglobulol, spathulenol, khusinol, khusinone, 12-ol (3.7–5.9 %); eremophila-1 (10),11-dien-
khusimone and khusinol acetate which contrib- 2α-ol (nootkatol) + ziza-6(13)-en-12-ol
uted to the characteristic odour of Vetiver oil. The (khusimol) (16.1–19.2 %); and eremophila-
oil also contained eugenol and isoeugenol. 1(10),7(11)-dien-2α-ol (isonootkatol) + (E)-
Among 94 isolated compounds, 21 aromatic vol- eremophila-1(10),7(11)-12-ol (isovalencenol)
atile components were identified from vetiver oil (5.6–6.9 %). The important compounds that
in China, accounting for 53.9 % of the total con- imparted the characteristic vetiver odour were
206 Poaceae
(Haiti, Texaroma) to 2.2 % (Java, Djasala) were octacosane; manool; hexadecanoic acid; nonadec-
the major components in all genotypes. Most ane; 1-dodecanamide, N,N-dimethyl-; o-guaiacol;
commercial oils contained 1.2–2.9 % vetivenic p-vinylguaiacol; (Z)-isoeugenol (tr) and decanal
acid except China, Guang Dong (25.2 %). (tr). In subsequent studies they found that tissue-
Nootkanone was found only in traces or absent cultured vs. natural plantlets of ‘Karnataka’ and
except for Java, Balansula (0.8 %). Also quantita- ‘Malaysia’ cultivars of vetiver grown in sterilised
tive differences were found between genotype soil resulted in the largest number of differences
oils in vetiselinenol, β-vetivone and α-vetivone. in compounds (Adams et al. 2008). The least
Adams et al. (2004) found that non-cleansed number of differences of compounds was between
(normal) vetiver had typical vetiver oil profile tissue-cultured (bacteria and fungi-free) vs. natu-
and yielded 0.02 % clear oil, whereas the tissue- ral plantlets grown in non-sterile soil. Fifty essen-
cultured (cleansed) vetiver yielded a 0.35 % yield tial oil components were compared, predominated
of light yellow oil containing large amounts of by khusimol (= zizanol) (18.92–25.68 %),
C19–C29 alkanes plus several alkanols along with epi-zizanone (3.20–6.81 %), (E)-isovalencenol
typical vetiver oil compounds, but lacked pre- (7.11–10.06 %), vetiselinenol (2.18–8.76 %),
sumed fungal metabolites such as β-funebrene, α-vetivone (1.76–3.50 %), α-cadinol (1.45–
prezizaene, α-amorphene and β-vetispirene. An 3.11 %), KI 1675 (unidentified sesquiterpene
unidentified biotic factor (apparently bacteria or alcohol, 2.79–15.10 %), KI 1625 (unidentified
fungi) appeared to enhance the oil production in sesquiterpene alcohol, 4.57–7.19 %), KI 1682
normal vetiver by both increasing yield and by (unidentified sesquiterpene alcohol, 1.49–3.35 %)
the generation of signature oil compounds. and KI 1698 (unidentified sesquiterpene alcohol,
Compounds found in the essential oil of 5-month- 2.18–3.14 %) as major components. The other
old non-cleansed vetiver plant included: khusi- fully identified minor components included: (E)-
mol (13.1–24.6 %), (E)-isovalencenol isoeugenol + prezizaene, khusimene, α-amorphene,
(8.9–11.8 %), sesquiterpene alcohol (M222) valencene, α-muurolene, δ-amorphene, elemol,
(2.5–4.5 %), α-cadinol (2–4.6 %), sesquiterpene β-vetivenene, trans-sesquisabinene hydrate,
alcohol (M220) (2.8–3.7 %), β-vetivone (2.1– khusimone, zizanal, nootkatol, 13-hydroxyl valen-
2.7 %), α-vetivone (1.5–2 %), vetiselinenol cene, α-costol, nootkatone, β-vetivone, cyclohexa-
(2.4–3 %), β-vetivenene (2–3.7 %) and other decanolide and hexadecanoic acid. For the
minor components: longicyclene, α-duprezianene, Malaysian genotype, zizanal was absent only in
β-elemene, β-funebrene, prezizaene, the ECHO (Ecological Concerns for Hunger
α-amorphene, β-vetispirene, 6,9-guaiadene, (E)- Organization) (natural) plants grown in sterilised
isoeugenol, khusimene, δ-amorphene, γ-vetivene, soil. For the Karnataka genotype, KI 1605 (uniden-
elemol, eudesm-7(11)-en-4-ol, nootkatol and tified sesquiterpene hydrocarbon) and KI 1679
14-hydroxy-δ-cadinene. Compounds found in (unidentified sesquiterpene alcohol) were present
the essential oil of 5-month-old cleansed vetiver in the tissue-cultured plants, but absent in normal
plant included: khusimol (11.5 %), (E)- (ECHO) plants.
isoeugenol (6.4 %), (E)-isovalencenol (4.2 %), Vetiver oil produced from root cultures of
tetracosane (5.2 %), pentacosane (4.9 %), trico- crown explants contained the highest content of
sane (4.3 %), hexacosane (3.8 %), epi-zizanone terpenoids (19.024 %) followed by rooted callus
(2.4 %), heptacosane (2.4 %), docosane (2.4 %) (10.35 %) and rootip explants (1.68 %) (Esyanti
and other minor components (<2 %): pentadec- et al. 2013). The following were indentified:
ane; hexadecane; sesquiterpene alcohol (M222); monoterpenes 4-ethyl-m-xylene, ψ-cumene,
α-cadinol; vetiselinenol; nootkatol; tetradecanol; 2-ethyl-1,4-dimethyl-benzene and dandurene and
heptadecanol; octadecane; sesquiterpene ketone sesquiterpene, phenol,2,4,6-tris (1-methylethyl).
(M218); sesquiterpene alcohol (M220); No meaningful differences were observed in
α-vetivone; β-vetivone; 14-hydroxy-δ-cadinene; the composition of vetiver essential oil from nine
khusimene; eicosane; heneicosane; nonacosane; geographical origins Brazil, China, Haiti, India,
208 Poaceae
Java, Madagascar, Mexico, Reunion and Salvador and 147.8 mg/g and cultivated in the beach were
(Champagnat et al. 2006). About 110 constitu- 321.7, 369.5 and 154.0 mg/g respectively.
ents were identified in the oils, mainly sesquiter- Contents of benzene–ethanol extractives were
penes. The characteristic constituents were 59.5 and 54.1 mg/g and ash were 81.7 and
β-vetispirene (1.6–4.5 %), khusimol (3.4– 71.7 mg/g between the two locations respec-
13.7 %), vetiselinenol (1.3–7.8 %) and α-vetivone tively. Among hydrolysis products in the leaf of
(2.5–6.3 %). Steam distillation of dried roots vetiver cultivated in experimental field and beach,
afforded a viscous light-brown oil in about 0.3– contents of glucose and xylose were higher with
1.0 % v/w yield with balsamic earthy and sweet 368.3 and 359.9 mg/g and 245.7 and 204.3 mg/g,
woody odour. Supercritical carbon dioxide respectively, and contents of arabinose, galactose
extraction resulted in high yield (3.2 %) of veti- and mannose were lower with total contents of
ver oil in significantly less time than hydrodistil- 58.6 and 55.8 mg/g, respectively.
lation (Martinez et al. 2004). Brazilian samples In vetiver shoots and leaves, the major com-
had a greater acid amount (especially zizanoic pounds found were 9-octadecenamide (33.50 %),
acid) and could be chemically transformed into 2,6,10,15,19,23-hexamethyl-2,6,10,14,18,22-
an alcohol (khusimol) of desirable sensorial tetracosahexaene (27.46 %) and
properties. Sensory evaluation indicated that the 1,2-benzendicarboxylic acid, diisooctyl ester
Brazilian volatile oil without acid could be used (18.29 %) (Huang et al. 2004). Three monoter-
in perfumery and the extract obtained with super- penes, two sesquiterpenes and one triterpene were
critical carbon dioxide could have application in found among the shoot volatiles. Nine compounds
food. Pripdeevech et al. (2010) found that com- cholesterol; 1,2-bis(4-hydroxy-3-methoxyphenyl)-
prehensive two‐dimensional gas chromatogra- propane-1,3-diol; 1-O-feruloylglycerol; 1-O-p-
phy–mass spectrometry was more efficient in coumaroylglycerol; trans-p-hydroxycinnamic
differentiating the quality of vetiver essential oils acid; vladinol E; vladinol F; tricin 4’-O-(erythro-
obtained from diverse extraction conditions in guaiacylglyceryl) ether, and tricin 4’-O-(threo-
terms of their volatile compositions and content guaiacylglyceryl) ether were isolated from aerial
than gas chromatography–mass spectrometry. parts of vetiver grass (Gao et al. 2012).
Vetiver alcohols (such as khusimol, isovalen- Studies showed that when vetiver grass was
cenol, vetiselinenol) and esters in Haitian vetiver exposed to moderate (20 % and 40 % polyethyl-
essential oils and vetiver acetates were purified ene glycol (PEG) 6000 solutions) and severe
and quantified using high-performance thin-layer (60 % PEG solution) water deficit for 6 days, the
chromatography by Paillat et al. (2012). High lin- plant injury degree (expressed as the parameters
ear correlation with R2 = 0.9995 for alcohols and of plant growth, cell membrane integrity, water
R2 = 0.9996 for acetates were obtained. Skeleton relations and photosynthesis) increased and con-
of vetiver alcohols were identified in both stan- tents of free and conjugated putrescine decreased
dard 1 reference mixtures RM1 (eudesmane, with the rise of PEG concentration (Zhou and Yu
helifolane) and standard 2 RM2 (cadinane, 2010). Under moderate water deficit, the plants
eremophilane, spirovetivane, zizaane/prezizaane could survive by the reduced osmotic potential
and cyclocopacamphane). and increased free and conjugated spermidine
and spermine in leaves. After subsequent rewa-
tering, the osmotic balance was re-established,
Aerial Part Phytochemicals most of the above investigated physiological
parameters were fully or partly reverted to con-
The contents of K, N, Ca, P, Mg and S in vetiver trol levels. The data indicated that vetiver grass
leaves were 11.2, 7.6, 4.3, 2.7, 2.8 and 1.5 mg/g, could cope well with the moderate water deficit/
respectively (Zhou and Yu 2012). Contents of drought stress by using the strategies of osmotic
cellulose, hemicellulose and lignin in vetiver leaf adjustment and maintenance of total contents of
cultivated in experimental field were 326.1, 380.2 free, conjugated and bound polyamines in leaves.
Chrysopogon zizanioides 209
Klebsiella pneumoniae (5, 10, 20 μg/ml), Recent studies showed that khusenic acid
Streptococcus faecalis (10, 20, 40 μg/ml), was 4 times more active than the standard drugs
Pseudomonas aeruginosa (10, 20, 40 μg/ml) and ciprofloxacin and nalidixic acid against the cip-
Salmonella typhi (20, 20, 40 μg/ml). The order of rofloxacin (CSC 101) and lomefloxacin
antifungal activity for the methanol, chloroform (LOMR5)-resistant Mycobacterium smegmatis
and hexane vetiver leaf extract based on mini- mutants, whereas khusimol was two times more
mum inhibitory concentration (MIC) values was active against the CSC 101 than nalidixic acid
as follows: Aspergillus niger (1.25, 5, 10 μg/ml), and ciprofloxacin (Dwivedi et al. 2013). Further,
Trichophyton mentagrophytes (5, 10, 20 μg/ml), the virulent strain H37Rv of Mycobacterium
Aspergillus flavus (5, 10, 20 μg/ml), Candida tuberculosis khusenic acid was two times more
albicans (10, 20, 20 μg/ml) and Saccharomyces active than nalidixic acid, while khusimol was
cerevisiae (10, 20, 40 μg/ml). The order of anti- equally active to nalidixic acid. In in-silico
bacterial activity for the methanol, chloroform docking study, khusenic acid showed better
and hexane vetiver root extract based on mini- binding affinity than khusimol with both sub-
mum inhibitory concentration (MIC) values was units of the bacterial DNA gyrase, which was
as follows: Staphylococcus aureus (1.25, 2.5 μg/ further confirmed from the in-vitro bacterial
ml), Escherichia coli (2.5, 5, 5 μg/ml), Klebsiella DNA gyrase inhibition study. Also, khusenic
pneumoniae (2.5, 5, 10 μg/ml), Pseudomonas acid and khusimol showed better intestinal
aeruginosa (2.50, 5, 10 μg/ml), Streptococcus absorption, aqueous solubility and ability to
faecalis (5, 10, 20 μg/ml) and Salmonella typhi penetrate the blood–brain barrier. Finally, khusi-
(5, 10, 20 μg/ml). The order of antifungal activity mol was found safe at the highest dose of
for the methanol, chloroform and hexane vetiver 2000 mg/kg body weight. Being edible, fragrant
root extract was as follows: Aspergillus niger natural products, khusenic acid and khusimol
(1.25, 5, 10 μg/ml), Trichophyton mentagro- from vetiver oil could have an advantage over
phytes (5, 10, 20 μg/ml), Aspergillus flavus (10, the existing synthetic drugs.
10, 20 μg/ml), Candida albicans (10, 20, 40 μg/
ml) and Saccharomyces cerevisiae (10, 20, 40 μg/
ml). Anticonvulsant Activity
Vetiver extracts were found to be more potent
against the gram negative Escherichia coli in Studies by Gupta et al. (2013) demonstrated that
comparison to Bacillus subtilis and Micrococcus oral administration of V. zizanioides extract
luteus, except for the alkaline methanol-soluble exerted significant anticonvulsant activity in mice
fraction which was more potent against M. luteus with an LD50 of 600 mg/kg body weight. Vetiver
and the methanol-soluble acid extract which was ethanol extract at a dose of 400 mg/kg signifi-
more active against B. subtilis (Prajna et al. cantly reduced flexion (l5.98–3.73 s), extension
2013). The water-soluble and methanol-soluble (13.73–0.96 s), clonus (14.07–4.93 s) and stupor
alkaline extract showed the lowest LC50 values (6.29–1.22 s) in the maximal electroshock stimu-
and hence the highest activity against E. coli. lation model. Further, it increases onset of clonic
Alkaline extract dissolved in methanol showed (88.25–708.32 s/30 min) and tonic (139.52–
the highest activity against B. subtilis and M. 1126.39 s/30 min) in the pentylenetetrazol
luteus. In contrast, the methanol-soluble alkaline model. Vetiver oil exhibited anticonvulsant activ-
fraction consistently showed better antifungal ity in both maximal electroshock (MES)- and
activity against Aspergillus niger and pentylenetetrazol (PTZ)-induced seizure models
Macrophomina phaseolina. The water-soluble in mice (Maignana Kumar et al. 2014). Vetiver oil
acidic fraction and the methanolic fraction were exhibited equivalent efficacy as that of sodium
more effective against Fusarium oxysporum. valproate in PTZ model and comparatively lesser
212 Poaceae
include: soil and water conservation, slope stabi- mal conductivity, the house is more comfortable
lisation, erosion control (road slopes and banks and energy saving. Moreover, both vetiver and
of reservoir), environmental protection (hedges clay are an abundant indigenous material, and
to keep off dust and heat), absorption of heavy construction using this composite is inexpensive
metals, water purification, disaster mitigation and and requires local-based skills, which can be con-
waste-water treatment. Non-conventional uses of sidered as labour-based appropriate technology
live vetiver plant include as forages for livestock (LBAT) for the community.
grazing, fish feed, ornaments (landscape plants,
hedge and potted plants for gardens, patios and
decks) and field boundary markers. Dried or par- Perfumery and Aromatherapy
tially dried plant (leaves, culms and roots) by-
products from non-processed plant materials are It is the major source of the well-known oil of
used as roof thatch, compost, mulch, potting vetiver, which is used in medicine and in perfum-
medium, mushroom medium (for shiitake and ery (Rao and Suseela 2000). Vetiver grass is also
oyster mushrooms), animal fodder and bouquet cultivated for the production of a commercially
displays; semi-processed products include handi- important essential oil used in perfumery and
craft, botanical pesticides, pots, low-cost silo, aromatherapy (Weyerstahl et al. 1996;
energy sources (ethanol, green fuel), furniture, Chowdhury et al. 2009). Vetiver oil and its con-
botanical pesticides and some industrial prod- stituents are used extensively for blending orien-
ucts. Fully processed products include essential tal type of perfumes and floral compounds, as
oil and its derived products, herbal medicine, well as in other cosmetic and aromatherapy
pulp and paper, fibreboard, pozzolana cement applications (Lavania 2003). Vetiver oil is a main
and industrial products, etc. ingredient in 36 % of all Western-quality per-
fumes and 20 % of all men’s fragrances. It is very
persistent and one of the finest fixatives known.
Construction Material Dried roots are used as sachets/stuffing material
to prepare ventilating screens that provide cool
VGA (vetiver grass ash) mortar was found to be air effect and pleasant aroma when moistened.
suitably adopted as a construction material for
foundations, marine structures, sewers and other
chemically exposed structures (Nimityongskul Erosion Control
et al. 2003). Test results revealed that the silica
content of vetiver grass ash was approximately The Vetiver system is the premier soil erosion
7 % higher and potassium oxide (K2O) content method outside of temperate zones (Anonymous
about seven times higher than in fly ash, and 2014). Narrow hedgerows of Vetiver grass will
according to ASTM requirement VGA could be spread out rainfall runoff across the slope, act as
classified as class C pozzolana. Paddy storage a filter to trap erosion sediment, create natural
silo could be made from vetiver grass and clay. terraces and reduce the velocity of rainfall runoff.
Studies by Hengsadeekul and Nimityongskul It is used on farm soil and for water conservation,
(2004) found vetiver–grass–clay composite pro- rehabilitation of eroded lands and prevention of
vided good insulation material that could reduce erosion on sloping lands. The Vetiver system
the effects of temperature and relative humidity when applied to such slopes significantly reduces
outside of the silos and extend the storage period the probability of land slippage and reduces the
and maintain the quality of paddy. Vetiver grass need for ‘hard solutions’. Applications include
could be used with clay to substitute bricks for highway, railway, riverbanks, public utility right
housing construction (Hengsadeekul and of ways, canal, dikes and levee slopes.
Nimityongskul 2005). Due to the prefabricated In north Queensland, the establishment of vet-
vetiver–clay block wall having a rather low ther- iver grass has been shown to control channel
Chrysopogon zizanioides 215
bank erosion, lower frequency of drain mainte- (khus) after oil extraction, root of Rosa dama-
nance, trap sediments in runoff water and reduce scena (rose) and bark of Eucalyptus citriodora
acidic loading by exposing less acid sulphate soil afforded biocoals with good calorific values
in the drain wall to oxidisation and leaching (Yadav et al. 2013). Owing to their similar fuel
(Troung et al. 2003). properties as high sulphur sub-bituminous coal,
they could be good candidates for cofiring.
Blending of these biocoals with high sulphur
Biofuel/Bioethanol/Biocoal coals will serve a dual purpose: (1) alternate fuel
Production and (2) reduction in SO2 emission.
to the unplanted control. Thus, there was no evi- The purifying effects of Vetiver to heavy metals,
dence of vetiver enhancing the biodegradation of N and P from pig farms were ranked as
crude oil in soil under the conditions of the trial. Zn > Cu > As > N > P > Pb > Hg. The Vetiver sys-
However, uses of vetiver grass in relation to tem has been identified to be an efficient, sustain-
petroleum-contaminated soils are promising for able and cost-effective system for effluent and
amelioration of slightly polluted sites, to allow solid-waste product treatment (Smeal et al.
other species to get established and for erosion 2003). Computer modelling output based on an
control. Glass house studies demonstrated that assumed maximum annual effluent output of 475
vetiver grass may promote the biodegradation of ML/year, and N concentration of 300 mg/L and P
benzo[a]pyrene (B[a]P) under flooded conditions of 1 mg/L, indicated that among the three grasses,
by plant roots by stimulating the microbial bio- vetiver required the least land for sustainable irri-
mass (Li et al. 2006). Microbial biomass was the gation in both N and effluent volume: 72.5, 104
main factor affecting dissipation of B[a]P under and 153 ha for vetiver, Kikuyu (Pennisetum clan-
flooded conditions. destinum) and Rhodes grass (Chloris gayana)
Liao et al. (2005) found that the below-ground respectively. The study of Hart et al. (2003)
biomass of V. zizanioides was greater than that of reported on the efficacy of Vetiver growing under
Cyperus alternifolius. In contrast, the above- hydroponic conditions to treat motel effluent,
ground biomass of C. alternifolius was greater which has been primary treated in septic tanks
than that of V. zizanioides. The annual biomass and to surface irrigate the motel gardens with the
yield of C. alternifolius (3406.47 g/m2) was treated effluent in a sustainable way. The best
2.3 times higher than that of V. zizanioides method trialled was effluent flow at 20 L/min
(1483.88 g/m2). The removals of N, P, Cu and Zn through Vetiver roots, a method that was highly
by harvesting vegetation were 4–7 times higher successful in reducing nitrogen concentrations.
in Cyperus alternifolius wetland than in V. zizani- The vetiver grass system to treat sewerage efflu-
oides wetland. ent on wetlands in southeast Queensland had
As a species of land reclamation, vetiver grass already shown itself to be a suitable alternative to
possesses various kinds of superior characteris- more expensive solutions to upgrade existing
tics and functions, such as rapid growth, huge sewage treatment plants (Ash and Truong 2003).
biomass, massive and long roots, strong abilities Singh et al. (2008) found in hydroponic studies
to control erosion and stabilise slopes and huge that Vetiveria zizanioides had potential for phy-
capacities for phytoremediation (Xia and Shu toremediation of phenol.
2003). Applications around the globe as well as The amount of arsenic removal was higher in
in China of vetiver grass include reclamation of Vetiveria zizanioides (Surat Thani ecotype) than
barren mountains or hills, contaminated water Vetiveria nemoralis (Balansa, Prachuabkirikhan
and soil, garbage landfills, mined lands, quarries, ecotype) (Srisatit et al. 2003). Arsenic accumula-
etc. From their series of experiments, a three- tion was higher in the roots than in the leaves.
pronged strategy was suggested for an integrated Both ecotypes grew well in arsenic soils. The
vetiver technique (IVT) for phytoremediation of number of clumps and diameter of clumps were
heavy metal contamination: use of vetiver for higher for V. zizanioides than V. nemoralis. Free
phytostabilisation of heavy metals, use of vetiver water surface for tannery waste-water post-treat-
for phytoextraction of heavy metals and/or its ment with Surat Thani ecotype vetiver grass at
potential use with chemical chelators and use of 0.1 m water level was the most efficacious for
vetiver for phytofiltration of heavy metals (Shu chromium removal with 89.29 % efficiency.
and Xia 2003). They suggested that IVT could be Vetiver grass possesses special characteristics
used as an integrated technique for environmen- that make it a good choice for phytoremediation
tal management of mining activities. Vetiver had of heavy metals and organic wastes (Danh et al.
a very high capacity to purify waste water when 2009). Vetiver can accumulate heavy metals, par-
used as a vetiver bamboo float (Kong et al. 2003). ticularly lead (shoot 0.4 % and root 1 %) and zinc
Chrysopogon zizanioides 217
(shoot and root 1 %). The majority of heavy met- soils, a vertisol and a lixisol, from Burkina Faso
als are accumulated in roots thus suitable for phy- (Abaga et al. 2014). Six months after vetiver
tostabilisation and for phytoextraction with planting treatment, endosulfan was not detected
addition of chelating agents. Vetiver can also in soils indicating the effectiveness of vetiver in
absorb and promote biodegradation of organic promoting adsorption and the disappearance of
wastes (2,4,6-trinitroluene, phenol, ethidium bro- endosulfan in both soils.
mide, benzo[a]pyrene, atrazine). Studies by Datta Successful rehabilitation of mine tailings and
et al. (2013) showed that vetiver grass can be landfills with elevated levels of heavy metals in
used as an in-situ phytoremediation agent to Australia, China and South Africa indicates that
remove tetracycline from waste water. Studies by the Vetiver system should provide a powerful
Danh et al. (2011b) showed that vetiver grass phytoremedial tool for the attenuation of the mer-
could be used for revegetation of lead- cury pollution problem in Yolo and Lake counties
contaminated soils. They found that a level of by trapping and containing both the air- and
applied CaCO3 about half of the lead concentra- water-borne insoluble mercury at sources and by
tion in soils was sufficient to improve vetiver reducing the soluble fraction in acid mine drain-
growth and survival and accumulate high con- age (Truong 2003). Studies in Tanzania found
centrations of lead in the roots. Danh et al. Vetiver grass performed better than Phragmites
(2011a) showed that phytoremediation of Cu and mauritianus in removing pollutant in the waste-
Zn contaminated soils by vetiver could generate water treatment in horizontal subsurface flow
revenue from the commercialisation of vetiver constructed wetlands (Njau and Mlay 2003). For
essential oil extracts. Studies showed that inocu- instance, it was found from the treatment plant
lation of vetiver grass with a functional endo- that the organic removal BOD (biological oxy-
phytic bacterium, Achromobacter xylosoxidans gen demand) was on average 61.85 % and
F3B, enhanced its phytoremediation potential in 67.47 % and COD (chemical oxygen demand) of
the removal of aromatic pollutants (Ho et al. 37.9 % and 46.2 % by Phragmites mauritianus
2013). and Vetiver grass respectively. Studies in
Antiochi et al. (2007) found vetiver grass to be Guangzhou, China, found that vetiver grass con-
have good phytoremediation activity for heavy structed wetland has great potential in treatment
metals like lead and zinc but not chromium and of highly concentrated landfill leachate after pre-
copper. Andra et al. (2009a) found vetiver grass treatment to reduce its ammonium nitrogen con-
to have a mechanism of high Pb tolerance, sug- centration to 383 mg/L or even lower (Lin et al.
gesting its potential usefulness for the remedia- 2003). Vetiver grass was found to be highly toler-
tion of lead-contaminated residential sites. ant to adverse conditions such as acid mine leach-
Vetiver grass could accumulate up to 19,800 and ate, containing high concentrations of Zn, Mn,
3350 mg Pb/kg dry weight in root and shoot tis- Pb, Cd, Cu and sulphates and could be useful in
sues, respectively, in a hydroponics set-up. In the the design of full-scale constructed wetland for
presence of 15 mM/kg EDTA, vetiver accumu- treatment of acid mine leachates (Shu 2003).
lated 4460 and 480 mg Pb/kg dry root and shoot
tissue, respectively, that were 15- and 24-fold
higher compared to those in untreated controls Water Treatment
(Andra et al. 2009b). Paz-Alberto et al. (2007)
found that vetiver grass possessed many benefi- Studies by Kantawanichkul et al. (2013) showed
cial characteristics to uptake Pb from heavily that domestic waste water could be treated by
PB-contaminated soil. It can be used to phytore- vertical flow constructed wetland systems planted
mediate urban soil with various contaminations with Cyperus alternifolius (umbrella sedge) and
by planting these grasses in lawns and public Vetiveria zizanioides (Vetiver grass). The former
parks. Vetiver grass was found to be effective in was found to be more efficient than vetiver grass
the phytoremediation of endosulfan in two cotton in terms of removal efficiency of COD (chemical
218 Poaceae
oxygen demand) and nitrogen in terms of total synthetic capacity, fast growth rate and relatively
Kjeldahl nitrogen (TKN) was 76 and 65 % high contents of cellulose and hemicellulose in
respectively at 20 cm/d hydraulic loading rates leaf and to grow well in the marginal land and be
for umbrella sedge compared to only 67 and considered as a potential lignocellulosic energy
56 % for Vetiver grass. Studies showed that V. plant (Zhou and Yu 2012).
zizanioides cultivated on floating beds had a great
ability to remove total nitrogen, ammonium
nitrogen, total phosphorus, phosphates, COD and Production of Activated Carbon
BOD and had a significant effect on purifying
eutrophicated water (Si et al. 2003). Liao and Gas and liquid adsorption studies showed that
Luo (2002b) that Vetiveria zizanioides and large quantities of lignocellulosic residues
Cyperus alternifolius had very stable effects on derived from the industrial production of essen-
the removal of COD and BOD in waste water tial oil from vetiver grass roots could be used for
from pig farm. There was no significant differ- the production of activated carbon (Gaspard et al.
ence between the two in the removal of COD, 2007).
BOD and soluble solids. High removal rates of
ammonium nitrogen and phosphate sulphur from
piggery waste water were found for both species Animal Feed
in spring and significant removal of total nitrogen
in waste water by both species in autumn, while The leaves of vetiver are a useful by-product to
significant removal of total phosphorus in waste feed cattle, goats, sheep and horses (Truong et al.
water was obtained only in V. zizanioides con- 2008). It is also used as cattle fodder in Tanzania.
structed wetlands (Lia and Luo 2002a). Studies Studies by Liu et al. (2003) found that digestibili-
by Xiong et al. (2011) found that using an inte- ties of gross energy, dry matter, crude protein,
grated constructed treatment system (CTS) with ether extract, crude fibre, calcium, phosphorus
vetiver growing in vertical constructed wetland and nitrogen-free extract in Vetiver grass hay
and Coix lacryma-jobi in floating beds had excel- were 29.65 %, 46.09 %, 23.15 %, 28.79 %,
lent efficacy in removal of COD and phosphorus 46.44 %, 61.00 %, 66.60 % and 36.25 %, respec-
from secondary effluents. tively. 1 kg dry matter of Vetiver grass hay could
provide 1.47 Mcal digestible energy, 13.4 g
digestible crude protein and 4.17 g ether extract,
Carbon Sequesterisation which indicated that Vetiver grass is a promising
feed resource for Donghsan goats.
Vetiver grass, a non-invasive C4 grass with fast
growing tufted root system reaching 3 m in one
year, could be an ideal global candidate to facili- Insect Pest Control
tate long-term locking of atmospheric carbon
below plough layer with reduced chances of Khusitoneol, a keto alcohol from vetiver oil,
being recycled into the atmosphere and replenish exhibited considerable juvenile hormone activity
soil carbon sink (Lavania and Lavania 2009). It against Lipaphis erysmi (mustard aphid) (Kalsi
afforded a holding potential of 1 kg atmospheric et al. 1985a). Of eight essential oils (vetiver grass,
carbon sequestered annually deep in the soils cassia leaf, clove bud, cedarwood, Eucalyptus
pool from one m2 surface area. Strategic estab- globulus, Eucalyptus citriodora, lemongrass and
lishment of vetiver in crop field, tree line, river, geranium) evaluated against the Formosan sub-
road and rail line embankments as hedge rows terranean termite, Coptotermes formosanus, veti-
could potentially contribute to carbon sequester- ver oil proved the most effective repellent because
ing and as a source of biomass and bioenergy. V. of its long-lasting activity (Zhu et al. 2001a).
zizanioides was found to have a very high photo- Nootkatone, a sesquiterpene ketone, isolated
Chrysopogon zizanioides 219
from vetiver oil was found to be a strong repel- ling at concentration as low as 5 μg/g sand and
lent and toxicant to Formosan subterranean ter- entirely inhibited termites’ tunnelling and paper
mite Coptotermes formosanus (Zhu et al. 2001b). consumption at concentrations higher than
The lowest effective concentration tested was 25 μg/g sand. Two-choice preference bioassays
10 μg/g substrate. Among eight valencenoid and stem borer larval survival experiments were
derivatives evaluated for their repelling activity conducted to determine the suitability of vetiver,
against Formosan subterranean termites, napier grass (Pennisetum purpureum) and maize
Coptotermes formosanus, for survival of stem borer, Chilo partellus (Van
1,10-dihydronootkatone was the strongest repel- den Burg et al. 2003). Results indicated that veti-
lent, and valencene was the weakest (Zhu et al. ver grass was highly preferred for oviposition but
2003). Maistrello et al. (2001b) found that sand that larval survival on vetiver grass was extremely
treated with vetiver oil or nootkatone at 100 μg/g low. Thus, vetiver has potential as trap crop com-
disrupted termite tunnelling behaviour. As a con- ponent of an overall ‘push–pull’ strategy to con-
sequence, after 21 days, wood consumption and centrate C. partellus oviposition away from the
termite survival were significantly lower com- maize crop and reduce subsequent population
pared with cedrene-treated or untreated sand development. This technology may also have
treatments. Maistrello et al. (2001a) found that application in rice stem-borer pest management.
nootkatone acted as a feeding deterrent, inducing
starvation that resulted in a complete loss of the
termite symbiont protozoa Pseudotrichonympha Landscaping
grassii, the most important flagellate species of
cellulosic digestion in Coptotermes formosanus. The Vetiver system can be applied for urban land-
In subsequent studies, they found that Vetiver scaping including beautification, slope stabilisa-
oil and especially its constituent nootkatone tion, traffic dividers, demarcation of walkways
affected Formosan subterranean termites and and prevention of urban erosion (Anonymous
their protozoa, acting as arrestants, repellents and 2014). Vetiver can make a city greener as a road
feeding deterrents, and represent a promising boundary, at roundabouts and in landscaping
natural alternative for the control of this invasive parks and resorts. It can make a city cleaner
pest (Maistrello et al. 2003). Ibrahim et al. (2004) through (1) its pond embankment filtration; vari-
reported that in sand barrier assays, a concentra- ous techniques in domestic, industrial and agri-
tion of 100 ppm of nootkatone and two of its cultural waste-water treatments; and water
derivatives, 1,10-dihydronootkatone and tetrahy- purification; (2) rehabilitation of contaminated or
dronootkatone, significantly reduced polluted water with the treatment of eutrophi-
Coptotermes formosanus survival, tunnel build- cated water, removal of effluents, heavy metals
ing and food consumption after a 12-day expo- and toxic substances; (3) treatments of landfills
sure. Termites preexposed to 100 ppm and garbage dumps with the removal of agro-
nootkatone-treated sand and placed in containers chemicals and pesticides and absorption of heavy
without nootkatone for 15 days continued to metals—a new approach of phytoremediation—
exhibit abnormal feeding and digging behav- and (4) dust reduction (Chomchalow 2012).
iours; survivorship, tunnelling and feeding It can make a city cooler using its evapo-
activities were significantly reduced by 83.5, transpiration function, heat reduction and the
63.2 and 95.4 %, respectively. cooling and refreshing effects of its dried massive
Vetiver oil was found to possess strong ter- roots. It can make a city safer serving as a wind-
miticidal activity against the Formosan subterra- break and utilising its ability to stabilise slopes of
nean termite, Coptotermes formosanus (Chen the road, riverbanks, ponds and shorelines. And,
et al. 2003). Vetiver oil decreased termite tunnel- finally, being a grass with a beautiful form and
220 Poaceae
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Eleusine indica
Agropyron geminatum Schult. & Schult.f., Bullgrass, Crabgrass, Crow’s Foot, Crowfoot
Chloris repens Steud. (inv.), Cynodon indicus Grass, Crowsfoot Grass, Dog Grass, Foul Foot,
(L.) Raspail, Cynosurus ara Buch.-Ham. ex Wall. Fowl Foot Grass, Fowl-Foot Grass, Goose Grass,
(inv.), Cynosurus indicus L., Cynosurus pectina- Goosegrass, India Goosegrass, Indian Goose
tus Lam., Eleusine distachya Trin. ex Steud. Grass, Indian Goosegrass, Iron Grass, Silver
(inv.), Eleusine distans Moench (illeg.), Eleusine Crabgrass, Wire Grass, Wire Grass, Yard Grass,
distans Link., Eleusine domingensis Sieber ex Yardgrass
Schult. (inv.), Eleusine glabra Schumach.,
Eleusine gonantha Schrank, Eleusine gouinii
E.Fourn., Eleusine inaequalis E.Fourn., Eleusine Vernacular Names
indica subsp. indica, Eleusine indica var. major
E.Fourn., Eleusine indica var. monostachya Arabic: Kalassindra (Chad)
F.M.Bailey, Eleusine indica var. oligostachya Bangladesh: Binna Chall, Chapra, Gaicha,
Honda, Eleusine japonica Steud., Eleusine mac- Malangakuri
rosperma Stokes, Eleusine marginata Lindl., Benin: Gamatori, Gomateri (Bariba), Tchouan
Eleusine polydactyla Steud., Eleusine rigidifolia (Berba), Torohundo (Yom)
E.Fourn., Eleusine scabra E.Fourn., Eleusine tex- Brazil: Ca-A Pi-1, Capim Criador, Capim Pe Da
tilis Welw. (inv.), Juncus loureiroana Schult. & Galinha, Capim-Da-Cidade, Capim-De-
Schult.f., Leptochloa pectinata (Lam.) Kunth, Burro, Grama-De-Coradoura, Grama-De-
Paspalum dissectum Kniph. (illeg.), Poa spicata Coradouro, Grama De Sapo (Portuguese)
Willd. ex Steud. (inv.), Triticum geminatum Burmese: Myet-Thakwa, Se-Gwa, Sin Ngo Let
Spreng. Kya, Sin Ngo Myet
South Africa: Indiese Osgras, Jongos Gras Edible Plant Parts and Uses
Spanish: Eleusine, Grama De Caballo, Grama De
Orqueta, Grama Dulce, Guarataro, Olotillo, The small seed is used as famine food and it can
Hierba Dulce, Natajo Dulce, Pata De Gallina, be cooked whole or ground into a flour and
Pata De Gallo, Pata De Ganso, Yerba De used in making cakes, gruels and alcoholic bev-
Camino, Yerba Dulce erages (Kunkel 1984; Facciola 1990; Harris
Swedish: Gåshirs 1995). Young plants are eaten raw or cooked
Tahitian: Tamamau, Tamaomao and used as a side dish with rice (Uphof 1968;
Taiwan: Nui Chin Tsao Tanaka 1976; Kunkel 1984; Facciola 1990).
Thailand: Yaa Teen-Ka Roots are edible raw (Kunkel 1984; Facciola
Tongan: Mohuku Siamane, Takataka, ‘A Leala, 1990).
A Le Ala
Togo: Adon’doulé, Tchama (Kabiyé)
Tongarevan: Mauku Botany
Uganda: Kasbanti
Ulithian: Fathil Eleusine indica is branching, erect or prostrate
Vietnam: Cỏ Mần Trầu, Tết Suất Thảo, Ngưu annual or short-lived perennial, and its culms are
Cần Thảo, Cỏ Vườn Trầu, Màng Trầu, Thanh tuft forming and branching from the base
Tâm Thảo, Cỏ Chỉ Tía, Ngưu Cân Thảo, Hang (Plate 3). Leaves are narrow and arranged in two
Ma (Tày), Co Nhả Hút (Thái), Hìa Xú Xan rows; the blade is glabrous, and sheath is hairy.
(Dao), Cao Day (Ba Na), Hất T’rớ Lạy (K’ho), Leaf blades are flat or folded, 15–30 cm long and
R’day (H’dong) 4–6 mm wide. Inflorescence is in a whorl of 2–7
Zambia: Rapoka (usually 5) digitate spikes from apex of culm,
Zimbawe: Mu Kha with single spikelet or two spikelets separated
below (Plate 4). Spikelets are 3–5 flowered, up to
4 mm long, awnless and dark green. Glumes are
membranous, with the lower part which is
Origin/Distribution 1–1.5 mm long, pointed, rough on the keel and
one-nerved and the upper part 3 mm long, sharply
The plant is indigenous to Africa, but long natu- pointed or tapering gradually to a point, with
ralised elsewhere, including South America, smooth keel, one- to five-nerved; lemmas are
Asia, Micronesia, American Samoa and most of similar in texture and shape to the glumes, ovate
the rest of the Pacific Islands from the tropical to and sharply pointed; palea is somewhat rough on
subtemperate regions. the keels. Grain is reddish brown to black,
oblong-ovate, ridged and striated.
Agroecology
Nutritive/Medicinal Properties
It is a major invasive weed of disturbed places,
irrigated fields and canals, including cultivated Twenty phenolic compounds were detected in
crops, pastures, gardens and roadsides, and also the leaves of Eleusine coracana subsp. coracana,
occurs in plantations and nurseries in the tropics E. coracana subsp. africana, Eleusine indica,
and subtropics (Plates 1 and 2). It is found from E. multiflora, E. tristachya, E. floccifolia and E.
near sea level to 2000-m elevation. It grows best compressa (Hilu et al. 1978). The leaf flavonoids
in moist, fertile, cultivated soil in full sunlight. It identified were orientin, isoorientin, vitexin,
is prevalent in disturbed areas, especially in isovitexin, saponarin, violanthin, lucenin-1 and
sandy soil. It is quick growing, long lived and tricin. The morphologically well-defined E. cora-
partial to wetter locations. cana–africana–indica group formed a unit in
Eleusine indica 231
Antidiabetic Activity
Cameroon, the whole plant is used for haemor- treat menorrhagia. A decoction of the plant is
rhagic cough (Sandberg et al. 2005). In Nigeria, used as cardiotonic. In Central African Republic
the plant is used as an anti-inflammatory and for (Oubangui), pounded plant is used as a bath for
convulsion in children (Obute 2005). Leaves of malaria in children (Vergiat 1969).
Tetracera alnifolia and Eleusine indica are
pounded and the poultice rubbed on skin for sca-
bies and skin eruption in Sierra Leone (Lebbie Other Uses
and Guries 1995). Eleusine indica leaves are
macerated and the infusion is employed for treat- The stems are used to make mats, baskets and
ing urine retention. Whole plant of Olax subscor- hats. The plant is suitable for paper manufacture.
pioidea + Eleusine indica + Eragrostis cilianensis The plant is sued in magic rituals where Malays
is made into semi-powder and is burned to fumi- hold the grain in their hand in spirit-summoning
gate non-violent mental patient by Gwandara rituals. In Bontoc, Philippines, the plant is used
tribe of Sabo Wuse in Niger state (Ibrahim et al. in mangmang rituals. Its strong roots make it a
2007). In Ghana, district of Gosomtwi-Atwima- good plant to prevent soil erosion. The plant is
Kwanwoma, a poultice of aerial parts of Eleusine used for silage and hay.
indica, is used to treat new and old wounds Crabgrass or crowfoot grass is considered in
(Agyare et al. 2009). In Central Nigeria (Kwara some regions to be a good fodder grass.
state), Eleusine indica leaves are crushed and Eleusine indica, Cynodon dactylon, Cyperus
mixed with shea butter (Butyrospermum para- rotundus and Equisetum ramosissimum showed
doxum) to make an ointment used to massage the promising potential for phytoremediation of heavy
painful parts of the body (Bhat et al. 1990). In metal pollution soil (Anh et al. 2011). The plants
Benin, E. indica roots are employed to treat diar- accumulated very high Pb (0.15–0.65 %) and Zn
rhoea; pulped roots are used locally for fractures (0.22–1.56 %) concentration in their roots.
and E. indica roots and Aframomum melegueta
are used for treating antalgic (Adjanohoun et al.
1989). A decoction of E. indica plant is used as a Comments
wash for boils and abscesses. An aqueous decoc-
tion of leaves of Commelina benghalensis, E. indica is ranked the 5th worst weed in the
Eleusine indica and Cleome viscosa is taken world and in Southeast Asia and an important
orally for treating female infertility. In the weed in over 60 countries (Holm et al. 1977).
People’s Republic of the Congo (Brazzaville),
aqueous decoction of the whole plant is sued for
treating urogenital infections (Adjanohoun et al. Selected References
1988). The same decoction is taken orally as a
mouth wash for influenza. In Togo, leaves and Adjanohoun EJ, Ahyi MRA, Aké Assi L, Akpagana K,
Chibon P, El-Adji A, Eymé J, Garba M, Gassita JN,
shoot of E. indica and ripe fruit of Aframomum
Gbeassor M, Goudote E, Guinko S, Hodouto KK,
melegueta are powdered diluted in lemon juice Houngnon P, Keita A, Keoula Y, Hodouto WP, Issa Lo
and taken orally for tachycardia (cardiac pain), A, Siamevi KM, Taffame KK (1986) Contributions
and aqueous decoction of the roots is used taken aux Études Ethnobotaniques et Floristiques au Togo.
Médecine Traditionelle et Pharmacopée Agence de
orally to treat urethral infection (Adjanohoun
Coopération Culturelle et Technique, Paris, 671 pp
et al. 1986). Pounded seeds of Aframomum Adjanohoun EJ, Ahyi AMR, Aké Assi L, Baniakina J,
melegueta and E. indica roots are taken orally for Chibon P, Cusset G, Doulou V, Enzanza A, Eymé J,
fever and convulsion. In Côte d’Ivoire, E. indica Goudoté E, Keita A, Mbemba C, Mollet J,
Moutsamboté J-M, Mpati J, Sita P (1988) Contribution
sap is applied on cuts, wound and injuries and is
aux Études Ethnobotaniques et Floristiques en
used for headache, for rib pain and as nose drop République Populaire du Congo. Médecine
(Bouquet and Debray 1974). Pulped roots are Traditionelle et Pharmacopée Agence de Coopération
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Morinda officinalis
Synonyms
Rubiaceae
Agroecology
cooking the roots of Morinda officinalis and with five anthraquinones (tectoquinone, alizarin
Pueraria with pumpkin, ginger, leek, salt and 1-methyl ether, lucidin ω-methyl ether,
pepper in a stew. In South China, Hong Kong and 1-hydroxyl-2,3-dimethylanthraquinone and
Macao, this plant has been developed into vari- 1 -hydroxy- 3 -hydroxymethylanthraquinone),
ous health foods, such as ‘Ba-ji-tian wine’, four iridoid glucosides (asperuloside, asperulo-
‘Ba-ji-zi-bu Gao’ (Li et al. 2009). sidic acid, desacetyl asperulosidic acid and
monotropein), a monoterpene glycoside (l-
borneol 6-O-β-D-apiosyl-β-D-glucoside), two
Botany sterols (β-sitosterol and oxositosterol), an ursane-
type triterpene (rotungenic acid) and a lactone
A perennial slender climbing shrub, several m compound (4R,5S)-5-hydroxyhexan-4-olide
long with greyish bark and angular twigs. Roots (Yoshikawa et al. 1995); succinic acid, nystose,
cylindrical 0.5–2 cm diameter, with longitudinal 1F-fructofuranosylnystose, inulin-type hexasac-
wrinkles and transverse cracks, dark grey or yel- charide and an heptasaccharide I (Cui et al.
lowish grey (Plate 1). Young shoots are violet 1995); 3-hydroxy-1,2-dimethoxy-anthraquinone
when young, tomentose becoming glabrous. (Xu et al. 2009); five anthraquinones including
Leaves on short petioles are opposite, simple, alizarin-1-methylether; 1,2-dimethoxy-3-hydro-
entire, lanceolate to obovate, 6–14 cm long by xyanthraquinone; 1-hydroxy-3-hydroxymethyl-
2.5–6 cm wide, base obtuse or rounded, apex anthraquinone; rubiadin-1-methylether and
acuminate, glossy dark green above, pale violet anthragallol-2-methylether (Zhu et al. 2009a);
and hairy on both sides especially the lower sur- 2-hydroxy-1-methoxy-anthraquinone monohy-
face below, stipule tubular thin, compressed, drate (Liu and Jiao 2009); physicion, rubiadin-
petiole short. Inflorescence in terminal umbellate 1-methyl ether, 2-hydroxy-1-methoxy-
panicles; 0.3–1.5 cm long, flowers are small, anthraquinone, 1,2-dihydroxy-3-methylanthra-
white to pale yellowish, calyx tubular-cyathiform quinone, 1,3,8-trihydroxy-2-methoxy-anthraquinone,
with unequal sepals, corolla tube short. Fruit is 2 -hydroxymethyl- 3 -hydroxyanthraquinone,
globose, drupe, scarlet when ripe and two seeded. 2-methoxyanthraquinone and scopoletin (Wu
et al. 2009); four anthraquinones 2-hydroxy-3-
hydroxymethylnthraquinone, 2-hydroxy-1-
Nutritive/Medicinal Properties methoxy-anthranquinone, rubiadin-1-methyl
ether, rubiadin (Bao et al. 2010); three anthraqui-
Morinda officinalis roots were found to contain: nones, 1,2-dimethoxy anthraquinone, alizarin-2-
anthraquinones, iridoids, oligosaccharides, poly- methyl ether and rubiadin-1-methyl ether (Liu
saccharides (Wu et al. 2009, 2013; Choi et al. et al. 2012); four anthraquinone compounds,
2005; Zhu et al. 2008, 2009a, b); organic namely, 2-hydroxy-3-hydroxymethyl-anthraqui-
acids, sugars, resins, vitamin C, an essential none, 2-hydroxy-1-methoxy-anthraquinone,
oil (NIMM 1999), anthraglucosides rubiadin-1-methyl ether and rubiadin (Wu et al.
(1-hydroxyanthraquinone) and phytosterols, 2013). Morinda officinalis root was found to con-
β-sitosterol, 2-methyl-anthraquinone, rubiadin- tain 89.5 % oligosaccharides and 0.19 % proteins
1-methyl ether and 24-ethylcholesterol (Li et al. and 12 mineral elements including Zn, Fe, Ca, K,
1991); eight anthraquinones rubiadin; rubiadin- P and Na but did not contain As, Cd and Pb (Li
1-methyl ether; 1-hydroxyanthraquinone; et al. 2008).
1-hydroxy-2-methylanthraquinone; 1,6-dihydrox Anthraquinones in M. officinalis root were
y-2,4-dimethoxyanthraquinone; 1,6-dihydroxy- found to be distributed in parenchymatous cells,
2-methoxyanthraquinone; 1-hydroxy-2- and the content of anthraquinones in the root
methoxyanthraquinone and physcion (Yang et al. gradually increased with plant age (Yao et al. 2004).
1992); an iridoid lactone, morindolide, and an The iridoid glycoside monotropein and
iridoid glucoside, morofficinaloside, together deacetylasperulosidic acid were isolated from the
Morinda officinalis 239
roots (Choi et al. 2005). Seven anthraquinones traose and inulopentaose (Feng et al. 2012). Five
and one coumarin were isolated upon further oligosaccharides were found in M. officinalis
fraction of the ethanol root extract; these com- root: 2.128–21.28 μg sucrose, 1.864–18.64 μg
pounds were identified as physcion, rubiadin-1- 1-kestose, 1.92–19.2 μg nystose, 1.912–19. 12 μg
methyl ether, 2-hydroxy-1- 1F-fructofuranosylnystose and 2.368–23.68 μg
methoxyanthraquinone, 1,2-dihydroxy-3-methyl bajijiasu (Deng et al. 2012). Inulin-type oligosac-
anthraquinone, 1,3,8-trihydroxy-2- charides with different DPs (degree of polymeri-
methoxyanthraquinone, 2-hydroxymethyl-3-hy- sations) were isolated from the roots with purities
droxyanthraquinone, 2-ethoxyanthraquinone and of >98 % (Yang et al. 2010). An acidic polysac-
scopoletin (Wu et al. 2009). charide APMO was isolated from Morinda offici-
Seventeen compounds were identified in M. nalis (Zhang et al. 2013). It predominantly
officinalis roots: physcion, 1-hydroxy-2-methyl- consisted of galacturonic acid, arabinose and
anthraquinone, 2-Hydroxy-1- galactose. Galacturonic acid was assigned to be
methoxyanthraquinone, rubiadin, rubiadin 1→4 glycosyl linkage in its skeleton. Seven inu-
1-methylether, 1,3-dihydroxy-2-methoxyanthra- lin-type oligosaccharides (DP = 3−9) were anal-
quinone, 3-hydroxy-2-methylanthraquinone, ysed using double-development performance
digiferruginol, 1,2-dimethoxy-3-hydroxyanthra- thin-layer chromatography and scanning densi-
quinone, 1,3-dihydroxy-2-hydroxymethyl- tometry in Morinda officinalis (Zhou et al. 2014).
anthraquinone, lucidin ω-ethyl ether, Thirty-four compounds representing 77.4 %
anthraquinone-2-carboxyllic acid, 7-hydroxy- of the volatile oil of M. officinalis root were iden-
6-methoxy-coumarin, fumaric acid, stigmasterol, tified (Liu et al. 2005). The main constituents
daucosterol and β-sitosterol (Zhang et al. 2010) (>2 %) were: borneol L 29.28 %, α-zingiberene
Polysaccharides of M. officinalis consisted 4.88 %, ar-curcumene 4.49 %,1-hexanol
mainly of glucose and fructose in the molar ratio 3.40 %,β-sesquiphellandrene 3.34 %, 2-amylfu-
of 1.29:2.71 (Zhu et al. 2009b). Six oligosaccha- ran 3.32 %, n-nonanal 2.17 %, L-camphor 2.07 %
rides were isolated from M. officinalis roots and and β-bisabolene 2.06 %. Forty-six volatile com-
identified as sucrose, inulin-type trisaccharide, ponents were identified from 15-year-old M. offi-
inulin-type hexasaccharide, inulotriose, inulote- cinalis roots, which accounted for 89.98 % in the
240 Rubiaceae
peak area of total ion chromatogram. Nineteen tective, renoprotective, anti-inflammatory, anti-
volatile components were identified from nociceptive, antitumour, anti-impotence and
10-year-old M. officinalis roots, which accounted antiosteoporotic attributes.
for 70.01 %. Fourteen volatile components were
identified from 6 to 8-year-old M. officinalis
roots, which accounted for 63.23 %. The volatile Antioxidant Activity
components extracted from M. officinalis root
mainly included acids, aldehydes, alcohols, Studies showed that a 10-day oral administration
alkenes, etc. Organic acids were the most abun- of M. officinalis extract reduced the fasting serum
dant, accounting for 65.28 % of volatile compo- glucose, hepatic and renal thiobarbituric acid
nents with palmitic acid accounting for 45.75 % reactive substances (TBARS) level and signifi-
followed by oleic acid. Twenty-six volatile com- cantly increased the hepatic superoxide dismutase
ponents were identified from M. officinalis, com- (SOD) and catalase (CAT) activities as well as
prising mainly of hexadecanoic acid (highest), glutathione (GSH) levels in streptozotocin-
followed by linoleic acid, oleic acid, diisobutyl induced diabetic rats (Soon and Tan 2002).
phthalate, 3-methyl-benzaldehyde and borneol Morinda officinalis extract was found to scavenge
(Yin et al. 2009). superoxide anion and hydroxyl radicals in the
The contents of monotropein from the differ- chemiluminescence reaction of luminol–H2O2–
ent processed products of M. officinalis root, CuSO4 system (Wu et al. 2006). M. officinalis
namely, crude drugs, woody part, salt processed root extract showed a 47.8 % 2,2-diphenyl-1-pic-
and conventionally processed, were found to be rylhydrazyl (DPPH) radical scavenging effect in
13.92 mg/g, 9.10 mg/g, 9.21 mg/g and TM3 Leydig cells with no significant cytotoxicity
12.86 mg/g, respectively (Xu et al. 2007). (Chang et al. 2008). The treatment of the cells
In traditional oriental medicine, the roots are with 250 μg/mL extract showed the most signifi-
reported to exert a tonic effect on kidneys, cant protective effect (64 %) in the cell viability
improving yang, especially for shengyangxu assay with a decreased lipid peroxidation level
(kidney yang deficiency) that causes impotence, (1.75 nmol/mg protein), increased testosterone
lack of libido, infertility, premature ejaculation, production (43.5 pg/mL), and improved superox-
frequent urination, incontinence, irregular men- ide dismutase (SOD) activity (7.49 units of SOD/
ses, cold and painful lower abdomen and chronic mg protein) and catalase (CAT) activity (74.6
inflammation of nerves (especially the sciatica units of CAT/mg protein). The findings indicated
nerves). They are useful in treating impotence, that M. officinalis root, as an antioxidant, pro-
spermatorrhoea, delayed menstruation, hyperten- tected functions of cultured mouse TM3 Leydig
sion, rheumatism, musculoskeletal atrophy, cells from H2O2-induced oxidative stress. Studies
arthritis and fatigue and ridding of damp cold. showed that M. officinalis polysaccharide supple-
Roots are described as having the following mentation resulted in (a) increased antioxidant
properties: antibacterial, hypotensive, adrenocor- enzyme activities and (b) decreased malondialde-
tical stimulant, antidepressant, aphrodisiac, hyde level in rats (Zhu et al. 2009b). An acidic
androgenic, cerebral restorative, urogenital polysaccharide APMO of M. officinalis exhibited
astringent, analgesic, musculoskeletal restor- excellent capability in scavenging DPPH radicals,
ative, antirheumatic and interferon inducent. chelating ferrous ions and inhibiting haemolysis
Scientific studies showed that the roots have of rats’ erythrocyte induced by hydrogen perox-
antioxidant, hypoglycaemic, hyperglycaemic, ide, which was stronger than those of vitamin C at
antidepressant, antifatigue, cardiovascular pro- high concentration (Zhang et al. 2013).
Morinda officinalis 241
and corticosterone; but did not reverse the levels with an increase in osteoclast cells. The findings
of alkaline phosphatase (ALP), TNF-α (tumour suggested that the root extracts may act as both a
necrosis factor-α) and interleukin IL-6 in serum suppressor of bone resorption and an enhancer of
of ovariectomised rats. The findings demon- bone formation in vivo and may have some
strated that the root extract reduced bone loss in favourable effects for preventing and treating the
ovariectomised rats, probably via the inhibition osteoporosis induced by sciatic neurectomy. The
of bone resorption, but was not involved with administration of M. officinalis polysaccharides
bone formation. Anthraquinones and polysaccha- (MOP) to osteoporotic rats induced by ovariec-
rides from Morinda officinalis could be respon- tomy resulted in an increase in bone mineral den-
sible for their antiosteoporotic activity. sity and mineral element concentration and a
Bioactivity-guided fractionation led to the decrease in serum cytokine level, indicating that
successful isolation of antiosteoporotic compo- MOP administration may play an important role
nents, i.e. physicion (1), rubiadin-1-methyl ether in thwarting development of osteoporosis (Zhu
(2), 2-hydroxy-1-methoxy-anthraquinone (3), et al. 2008).
1,2-dihydroxy-3-methylanthraquinone (4), Compounds 1,3,8-trihydroxy-2-methoxy-
1,3,8-trihydroxy-2-methoxy-anthraquinone (5), anthraquinone (1), 2-hydroxy-1-methoxy-anthra-
2-hydroxymethyl-3-hydroxyanthraquinone (6), quinone (2) and rubiadin (3) decreased the
2-methoxyanthraquinone (7) and scopoletin (8) formation of bone resorption pits, the number of
from an ethanolic extract of the roots of Morinda multinucleated osteoclasts and the activity of
officinalis (Wu et al. 2009). Among them, com- tartrate-resistant acid phosphatase (TRAP) and
pounds 2 and 3 promoted osteoblast prolifera- cathepsin K in the coculture system of osteo-
tion, while compounds 4 and 5 increased blasts and bone marrow cells in the presence of
osteoblast alkaline phosphatase activity. All of 1,25-dihydroxyvitamin D(3) and dexamethasone
the isolated compounds inhibited osteoclast (Bao et al. 2011). They also enhanced the
tartrate-resistant acid phosphatase (TRAP) activ- apoptosis of osteoclasts induced from bone mar-
ity and bone resorption, and the inhibitory effects row cells with M-CSF and RANKL. Further,
on osteoclastic bone resorption of compounds 1 they improved the ratio of mRNA and protein
and 5 were stronger than that of other com- expression of OPG and RANKL in osteoblasts,
pounds. The results suggested that M. officinalis interfered with the JNK and NF-κB signal path-
and its anthraquinones may have therapeutic way and reduced the expression of calcitonin
potential against osteoporosis. receptor (CTR) and carbonic anhydrase II (CA
Pharmacological studies showed that M. offi- II) in osteoclasts induced from bone marrow cells
cinalis root extract enhanced the expression of with M-CSF and RANKL. The findings indicated
core-binding factor α1 (cbfα1), a key transcrip- that the anthraquinone compounds from M. offi-
tion factor for osteoblast differentiation (Wang cinalis were potential inhibitors of bone resorp-
et al. 2004), and increased the proliferation, alka- tion and bone loss. Morinda officinalis capsules
line phosphatase activity and osteocalcin (Li at all doses (90, 270 and 810 mg/kg/day) were
et al. 2004a). Animal preventive and therapeutic able to significantly prevent the ovariectomised-
studies showed that M. officinalis aqueous root induced loss of bone mass due to diminishing
extract administered to sciatic neurectomised sig- serum tartrate-resistant acid phosphatase
nificantly and dose-dependently suppressed the (TRAP), and alkaline phosphatase (AKP), levels
decrease in hind limb thickness, tibia failure load, while elevating osteocalcin level in the plasma
bone mineral density and tibia Ca and P contents (Li et al. 2014). Morinda officinalis capsules also
with an increase in serum osteocalcin levels (Seo enhanced the bone strength and prevented the
et al. 2005). Further, the root extracts also signifi- deterioration of trabecular microarchitecture.
cantly and dose-dependently suppressed the The results suggested that Morinda officinalis
decrease in histomorphometrical parameters of capsules possessed potent antiosteoporotic activ-
the tibia such as volume, length and thickness of ity which could be an effective treatment for
trabecular bone and thickness of cortical bone postmenopausal osteoporosis.
244 Rubiaceae
Oral treatment of acutely stressed blood stasis as a tonifying and replenishing natural herb med-
rats with M. officinalis root ethanol extract (3, 6, icine could be used to enhance reproductive
12 g/kg bw/day) for 5 days eliminated blood functions and that Raman spectroscopy could be
stasis, inhibited rate of platelet aggregation and an applicable technology for screening active
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Eurycoma longifolia
Common/English Names
Origin/Distribution
Ali’s Umbrella, Ali’s Walking Stick, Tongkat
Ali, Long Jack, Longjack, Malaysian Ginseng, The species is found in Indonesia and Malaysia
Manotes Asiatica, Picroxylon Siamese and, to a lesser extent, Thailand, Vietnam and
Laos. In Indonesia, this species only occurs natu-
rally in Sumatra and Kalimantan.
Vernacular Names
Wood/Root Phytochemicals
Plate 4 (a) Trunk base and root with bark removed, (b) Tongkat ali processed product
isolated from E. longifolia (Itokawa et al. from E. longifolia roots (Ang et al. 2002).
1991b). New cytostatic quassinoids, Eurycomanone, 13,21-dihydroeurycomanone,
6α-hydroxyeurycomalactone, longilactone and eurycomanol, longilactone, 14,15β-
14,15β-dihydroxyklaineanone, were isolated dihydroxyklaineanone and eurycomanol-2-O-β-
from the wood of Eurycoma longifolia glucopyranoside were identified in the n-butanol
with three cytotoxic quassinoids, 11-dehy- fraction (Chan and Choo 2002). Three new
droklaineanone, eurycomalactone and [n-pentyl β-carboline-1-propionate, 5-hydroxy-
5,6-dehydroeurycomalactone, and with seven methyl-9-methoxycanthin-6-one and 1-hydroxy-
cytotoxic tirucallane-type triterpenes, niloticin, 9-methoxycanthin-6-one] and 19 known
dihydroniloticin, piscidinol A, bourjotinolone A, β-carboline alkaloids including 9-methoxycan-
3-episapelin A, melianone and hispidone thin-6-one and canthin-6-one were isolated from
(Itokawa et al. 1992). A new C19 quassinoid, E. longifolia roots (Kuo et al. 2003b). A new
6α-hydroxyeurycomalactone, and eurycomalac- C19-quassinoid-type glycoside named euryco-
tone were isolated from the roots (Chan et al. maoside was isolated from the roots (Bedir et al.
1992). Two new 1,2-seco-1-nor-6(5 → 10)abeo- 2003). Sixty-five compounds were isolated from
picrasan-2,5-olide skeleton quassinoids, eurylac- E. longifolia roots including four quassinoid
tones A and B, and two new C18 skeleton diterpenoids including new ones eurycomalide A,
quassinoids 3 and 4 were isolated from Eurycoma eurycomalide B, 13β,21-dihydroxyeurycomanol
longifiola together with two known C18 quassino- and 5α,14β,15β-trihydroxyklaineanone (Kuo
ids 5 and 6 (Itokawa et al. 1993a). A new 1, et al. 2004). A quassinoid compound,
2-seco-1-nor-6(5–10)-abeo-picrasan-2, 5-olide 4,5,7,8,17-penta-hydroxy-1 4,18-dimethyl-6-
skeleton quassinoid named eurylactone and two methylene-3,10-dioxapenta-cyclo-[9.8.0.0.0.0]
new C19- and C20-skeleton quassinoids were iso- nona-dec-14-ene-9,16-dione methanol solvate
lated from Eurycoma longifolia wood (Itokawa dihydrate, a natural eurycomanone, was isolated
et al. 1993b). One compound was found to have a from Eurycoma longifolia roots (Teh et al. 2009).
longilactone-type C19-skeleton, and the others a A novel 2,3-dehydro-4α-hydroxylongilactone
klaineanone-type13β,18-dihydroeurycomanol, and two known phenyl propanoids, 2,3-dihydroxy-
eurycomanol-2-O-β-d-gluco-pyranoside, euryco- 1-(4′-hydroxy-3′-methoxyphenyl)-propan-1-one
manol and eurycomanone were identified in and scopolin, were isolated from the roots
E. longifolia (Ang et al. 1995a, b). Eurycomanone (Teh et al. 2010). Five compounds were obtained
(1), longilactone (2), 14,15β-dihydroxyklaineanone from E. longifolia and identified as scopoletin,
(3), 15β-acetyl-14-hydroxyklaineanone (4), 6α- 9-methoxycanthin-6-one, 7-methoxy-β-carbolin-
hydroxyeurycomalactone (5) and eurycomalac- 1-propionic acid, laurycolactone A and
tone (7) were isolated together with β,12α,15β- 7-methoxyinfractin (Han et al. 2011). Five new
triacetyleurycomanone from Eurycoma longifolia quassinoids, eurylactone E, eurylactone F, eury-
(Chan et al. 1998). Eurycomanone was recovered lactone G, eurycomalide D and eurycomalide E,
with the highest concentration, being about 16.8– along with ten known quassinoids including
39.6-fold higher than the other quassinoids 2, 3, 5 13β-methyl,21-dihydroeurycomanone, were iso-
and 7 but 145.3-fold higher than 4 which showed lated from E. longifolia roots (Park et al. 2014).
the lowest concentration. Three novel quassino- The volatile compounds identified in the head-
ids, eurylactones A–C, were isolated from the space of E. longifolia extracts were as follows:
roots (Ang et al. 2000b). Three major canthi- curcumene; (R)-(−)-massoilactone; 3-phenoxy-
none alkaloids, 9-methoxycanthin-6-one, 1-propanol; octanoic acid; benzoic acid; acetic
3-methylcanthin-5,6-dione and its 9-methoxy acid; 2-phenoxyethanol; 4 ethynyl-4-hydroxy-
analogue, were detected in Eurycoma longifolia 3,5,5-trimethyl-2-cyclohex-1-enone; butanol, 3 –
(Choo and Chan 2002). Quassinoids, eurycolac- methyl; 1-butanol; 1-pentanol; 2-hexadecanol;
tone D, eurycolactone E, eurycolactone F, laury- acetol; nonanal; 2-methylhexanol, benzaldehyde;
colactone B and eurycomalactone were isolated 2,3-butanediol, [S-(R*,R*)]-; butyrolactone;
Eurycoma longifolia 255
Four canthin-6-one alkaloids, namely, cytotoxic effect (IC50 ≤ 20.0 μg/ml) against pelvic
9-methoxycanthin-6-one, 9-methoxycanthin-6- cancer (RD) and epidermoid carcinoma (KB) cell
one-N-oxide, 9-hydroxycanthin-6-one and lines; however, it did not show any significant
9-hydroxycanthin-6-one-N-oxide, and one quas- cytotoxic effect against prostate DU-145 cancer
sinoid, eurycomanone, isolated from E. longifo- cell line (IC50 = 33.0 mg/ml) (Nurhanan et al.
lia roots, exhibited cytotoxicity on human breast, 2002). Petroleum ether and aqueous extracts
colon, fibrosarcoma, lung, melanoma, KB and showed insignificant cytotoxic effect
murine lymphocytic leukaemia (P-388) cell lines (IC50 > 20.0 μg/ml) against all the cancer cell lines
(Kardono et al. 1991). Eurycomanone was inac- tested. No toxic effect was observed for these
tive against murine lymphocytic leukaemia extracts when treated on the normal Chang’s liver
(P-388) but was significantly active against the cell line. E. longifolia root extracts and fractions
human cell lines tested including KB-V1 (a exerted a direct antiproliferative activity on human
multidrug-resistant cell line derived from KB). breast cancer cell line MCF-7 (Tee and Azimahtol
The two β-carboline alkaloids β-carboline-1- 2005). Three active fractions, F5, F6 and F7, dis-
propionic acid and 7-methoxy-β-carboline-1- played IC50 values of 6.17 , 4.40 and 20 μg/ml,
propionic acid were not significantly active. respectively. The resultant from F7 purification,
Beta-carboline alkaloids 9-methoxycanthin- 6- F16, exhibited a higher cytotoxic activity towards
one and canthin-6-one isolated from the roots MCF-7 (IC50 = 15.23 μg/ml) and a certain degree
demonstrated significant cytotoxicity in-vitro of selectivity against a normal breast cell line
against human lung cancer (A-549) and MCF-10A (IC50 = 66.31 μg/ml). F16 significantly
human breast cancer (MCF-7) cell lines increased apoptosis in MCF-7 cells through the
(Kuo et al. 2003b). Quassinoids, 6α- modulation of Bcl-2 protein levels. In-vitro studies
hydroxyeurycomalactone, longilactone and 14,15β- showed that the F16 fraction from E. longifolia
dihydroxyklaineanone; 11-dehydroklaineanone, exerted antiproliferative action and growth inhibi-
eurycomalactone and 5,6-dehydroeurycomalac- tion on human breast cancer MCF-7 cells by apop-
tone; and seven cytotoxic tirucallane-type triter- tosis induction through Bcl-2 protein pathway and
penes, niloticin, dihydroniloticin, piscidinol A, was independent of caspase-9 and p53 pathways
bourjotinolone A, 3-episapelin A, melianone and (Tee et al. 2007). Among the tested quassinoid
hispidone isolated from the roots, exhibited compounds from E. longifolia, eurycomalactone
potent cytotoxic activity against murine lympho- displayed the most potent activity against all the
cytic leukaemia P388 and human KB cancer cells tested cell lines: murine colon carcinoma colon
(Iotakawa et al. 1992). 26-L5 (IC50 = 0.70 μM), murine B16–BL6 mela-
Eurycomalactone; 5,6-dehydroeurycomalatone; noma (IC50 = 0.59 μM), Lewis lung carcinoma
7α-hydroxyeurycolalactone; 13α(21)- LLC (IC50 = 0.78 μM) and human lung adenocarci-
epoxyeurycomanone;15-acetyl-13α(21)- noma A549 (IC50 = 0.73 μM) (Miyake et al. 2010a).
epoxyeurycomanone; 12,15-diacetyl-13α(21)- These activities were comparable to clinically
epoxyeurycomanone; 12-acetyl-13,21-dihydroeu- used anticancer agent doxorubicin (colon 26-L5,
rycomanone; 15β-acetyl-14-hydroxyklaineanone; IC50 = 0.76 μM; B16–BL6, IC50 = 0.86 μM; LLC,
6α-acetoxy-14,15β-dihydroxyklaineanone; and IC50 = 0.80 μM; A549, IC50 = 0.66 μM).
6α-acetoxy-15β-hydroxyklainenone, from E. lon- Jiwajinda et al. (2002) found the most active
gifolia leaves, exhibited moderate antileukemic quassinoid compound isolated from the leaf, for
against human P338 cell lines with IC50 values of inhibition of tumour promoter-induced Epstein–
14, 6.6, 7.2,0.94, 7.8, 12 and 15 μg/ml, respec- Barr virus activation (antitumour promotion) was
tively (Morita et al. 1993a). Eurycoma longifolia 14,15β-dihydroxyklaineanone (5, IC50 = 5 μM).
methanol extract was found to exhibit antiprolif- Of 65 compounds, isolated from the roots, com-
erative activity in-vitro against human HT-1080 pounds 12, 13, 17, 18, 36, 38, 59 and 62 demon-
fibrosarcoma cells (Ueda et al. 2002). The metha- strated strong cytotoxicity towards human lung
nolic E. longifolia root extract showed significant cancer (A-549) cell lines; however, 12, 13, 17,
Eurycoma longifolia 259
38, 57, 58 and 59 exhibited strong cytotoxicity lowest IC50 values of 24.9 μM, 11.8 μM and
towards human breast cancer (MCF-7) cell lines 44.1 μM, 14.1 μM towards MCF-7, MGC-803
(Kuo et al. 2004). Several quassinoid compounds cancer cell lines, respectively, while compound 6
11, 23 and 24 exhibited cytotoxicity towards the exhibited moderate cytotoxicity towards HT-29,
highly metastatic HT-1080 human fibrosarcoma MCF-7, LOVO, BGC-823, MGC-803, HepG2,
cell line and showed potent cytotoxicity (IC50 val- HeLa and A549 cancer cell lines (Meng et al. 2014).
ues 0.93–1.1 μM) (Miyake et al. 2009). Among A partially purified quassinoid (eurycoma-
the compounds isolated from E. longifolia stem, none, 13α(21)-epoxyeurycomanone and
9,10-dimethoxycanthin-6-one (IC50 = 5.0 μM), eurycomanol)-rich fraction (TAF273) of E. lon-
10-hydroxy-9-methoxycanthin-6-one gifolia root extract exhibited antiangiogenic
(IC50 = 7.2 μM), dihydroniloticin (IC50 = 8.2 μM) activity (Al-Salahi et al. 2013). It caused signifi-
and 14-deacetyleurylene (IC50 = 3.2 μM) dis- cant suppression in sprouting of microvessels in
played stronger cytotoxic activity against a rat aorta with IC50 11.5 μg/ml. TAF273 (50 μg/
HT-1080 human fibrosarcoma cell line than the ml) showed remarkable inhibition (63.13 %) of
positive control 5-FU (IC50 = 9.2 μM) (Miyake neovascularization in chorioallantoic membrane
et al. 2010b). of chick embryo. In-vitro, TAF273 significantly
Eurycomanone, from E. longifolia, was found inhibited the major angiogenesis steps such as
to be cytotoxic on cancerous cells (CaOv-3, proliferation, migration and differentiation of
HeLa, HepG2, HM3KO, MCF-7) by inducing human umbilical vein endothelial cells (HUVEC).
apoptosis through upregulation of p53 and Bax The results demonstrated that the antiangiogenic
and downregulation of Bcl-2 independently of activity of TAF273 may be due to its inhibitory
functional E6 and E6-AP activity (Mahfudh and effect on endothelial cell proliferation, differen-
Pihie 2008). The characteristics of apoptosis tiation and migration which could be attributed to
including chromatin condensation, DNA frag- the high content of quassinoids in E. longifolia.
mentation and apoptotic bodies following eury-
comanone treatment were observed.
Eurycomanone was less toxic on normal cells Antimalarial Activity
(MDBK, Vero). Similarly, Zakaria et al. (2009)
found that eurycomanone was cytotoxic on can- Eurycoma longifolia aqueous extract exhibited
cerous liver cell, HepG2 by inducing apoptosis antiplasmodial activity against chloroquine-
through the upregulation of p53 and Bax and resistant Plasmodium falciparum strain (W2)
downregulation of Bcl-2. It was less toxic on nor- with IC50 value ≤ 4 μg/ml (Hout et al. 2006).
mal cells Chang’s liver and WLR-68. Wong et al. Hydroxycanthin-6-one, eurycomalactone,
(2012) reported that eurycomanone inhibited eurycomanone and eurycomanol isolated from
A549 lung cancer cell proliferation in a dose- Eurycoma longifolia and 6-hydroxy-5,6-
dependent manner at concentrations ranging dehydroeurycomalactone exhibited in-vitro anti-
from 5 to 20 μg/ml with IC50 value of 5.1 μg/ml. malarial activities against a multidrug-resistant
Eurycomanone at viable therapeutic concentra- Thailand strain (K-1) of Plasmodium falciparum
tions of 5–20 μg/ml exhibited significant antipro- (Chan et al. 1986). The two β-carboline alkaloids
liferative and anti-clonogenic cell growth effects β-carboline-1-propionic acid and 7-methoxy-β-
on A549 lung cancer cells. The treatment also carboline-1-propionic acid, isolated from the
resulted in suppression of the lung cancer cell roots, demonstrated significant antimalarial
tumour markers (heterogeneous nuclear ribonu- activity as judged by studies conducted with
cleoprotein (hnRNP) A2/B1, p53 tumour sup- cultured Plasmodium falciparum strains
pressor protein) and several known cancer cell (Kardono et al. 1991). The antimalarial activity
growth-associated genes (prohibitin, annexin 1 of Eurycoma longifolia extract (containing
and endoplasmic reticulum protein 28). Cytotoxic 13β,18-dihydroeurycomanol, eurycomanol-2-
compounds 2 and 5 from the roots exhibited the O-β-d-gluco-pyranoside, eurycomanol and
260 Simaroubaceae
decreasing the refractory period between the dif- mals, but the E. longifolia-treated groups showed
ferent series of copulation (Ang and Sim 1997b). lower sexual performance in mounting, intromis-
Similar copulatory effects were observed on sion and ejaculation than the testosterone-treated
middle-aged male rats but the effect was much group (Ang et al. 2000a). Also, E. longifolia pro-
smaller (Ang and Lee 2002b). E. longifolia frac- moted the growth of both ventral prostate and
tions produced a dose-dependent, recurrent and seminal vesicles as compared with the control,
significant increase in the episodes of penile but the growth of sexual accessories at 800 mg/kg
erection when compared with control male rats; of butanol, methanol, water and chloroform frac-
400 mg/kg of chloroform, methanol, water and tions of E. longifolia Jack was less than that of
n-butanol extracts exhibited respective penile the testosterone-treated group. In further studies
erection indices of 28.32, 31.32, 39.25 and 36.49, 800 mg/kg of butanol, methanol, water and chlo-
while 800 mg/kg further increased the indices to roform fractions of E. longifolia significantly
32.00, 35.32, 40.24 and 45.29 (Ang and Sim increased the levator ani muscle to 58.56, 58.23,
1997a). However, none of the treated rats exhib- 60.21 and 62.35 mg/100-g body weight, respec-
ited any homosexual mountings during the 1-hour tively, when compared with the control
observation period. They also found that chronic (untreated) in the uncastrated intact male rats,
administration of Eurycoma longifolia exerted and 49.23, 52.23, 50.21 and 52.35 mg/100-g
aphrodisiac effect on sexually naive male mice as body weight, respectively, when compared to
shown by the slow and transient reduction in hes- control (untreated) in the testosterone-stimulated
itation time and also a similar manner in the castrated intact male rats (Ang and Cheang
increase in the % of sexually naive male mice 2001). Hence, the pro-androgenic effect as shown
scoring right choice throughout the investigation by this study further supported the traditional use
period (Ang and Sim 1998a). In another study, E. of this plant as an aphrodisiac. Separate studies
longifolia modified the orientation activities of showed that E. longifolia root extracts produced a
the treated male rats in that they significantly dis- dose-dependent, recurrent and significant
played more frequent and vigorous mounting, increase in the episodes of penile reflexes as evi-
licking and anogenital sniffing towards the recep- denced by increases in quick flips, long flips and
tive females, and it further intensified self- erections of the treated male rats during the
orientation as indicated by the increased 30-minute observation period (Ang et al. 2001).
grooming of the genitals compared to the con- The results provided further evidence that E. lon-
trols (Ang and Sim 1998c, 1999). In addition, gifolia increased the aphrodisiac potency activity
rats treated with 800 mg/kg of methanol, water in treated animals. Ang and Ngai (2001) found
and butanol extracts of E. longifolia continued to that fractions of E. longifolia decreased the hesi-
show confinement to a particular area of the cage tation time of noncopulator male rats, throughout
(around the female), thus showing restriction in the investigation period using an electrical age.
movement as compared to the controls. However, Separate studies showed that a high dose (800 mg/
the treated males possessed a lack of interest in kg) of E. longifolia methanol, chloroform, water
the external environment as indicated by a reduc- and butanol fractions could increase libido in
tion in exploration, raring and climbing on the middle-aged male rats by increasing mount fre-
cage wall. In sexually naïve male rats, E. longifo- quency (Ang and Lee 2002a). It was demon-
lia treatment enhanced sexual motivation as evi- strated that middle-aged male rats treated with
denced by the high level of both the total number 800 mg/kg of E. longifolia Jack increased orien-
of successful crossovers, mountings, intromis- tation activities towards the receptive females
sions and ejaculations during the 9–12th week (anogenital sniffing, licking and mounting),
observation period (Ang and Sim 1998b). Studies increased genital grooming towards themselves
with inexperienced castrated male rats showed and restricted movements to a particular area of
that E. longifolia produced a dose-dependent the cage but decreased interest in the external
increase in sexual performance of the treated ani- environment (climbing, raring, exploration) as
Eurycoma longifolia 263
compared with the controls during the investiga- agent for reversing the effects of oestrogen by
tion period (Ang and Lee 2002c). Treatment with increasing spermatogenesis, sperm counts and
various fractions of E. longifolia 0.5 g/kg sperm motility in estradiol-treated rats after four-
enhanced the sexual qualities of middle-aged teen consecutive days of treatment.
male rats decreasing their hesitation time as com- Oral treatment of rats with a standardised
pared to controls (3-ml/kg saline) (Ang et al. methanol extract of E. longifolia containing the
2003a). More than 50 % of the male rats scored major quassinoid constituents of 13α(21)-
right choice after 2 weeks post-treatment. They epoxyeurycomanone, eurycomanone, 13α,21-
found E. longifolia continued to enhance sexual dihydroeurycomanone and eurycomanol, at
motivation in adult, middle-aged male mice and doses of 50, 100 and 200 mg/kg, significantly
in retired breeders (Ang et al. 2003b). They found increased sperm count by 78.9, 94.3 and 99.2 %,
a transient increase in the percentage of male respectively, when compared with that of control
mice responding to the right choice after chronic (Chan et al. 2009).The low count, poor motility
consumption of the fractions with 50 percent of and abnormal morphology of the spermatozoa
the adult middle-aged male mice treated with E. induced by the Andrographis paniculata fraction
longifolia. were significantly reversed by the standardised E.
Consumption of E. longifolia aqueous extract longifolia MeOH extract. The plasma testoster-
enhanced both sexual behaviour and sperm qual- one level of the rats treated with the standardised
ity in rats (Mat Noor et al. 2004). The extract MeOH extract at 200 mg/kg was significantly
(150 mg/kg)-treated male rats exhibited the increased when compared with that of the control
shortest period of time to reach the female rats and infertile animals. Interestingly, eurycoma-
compared to other doses and control. Oral admin- none alone was detected in the rat testis homog-
istration of the aqueous extract resulted in enates and may have contributed towards the
increased sperm count, viability and motility. improvement of sperm quality. Thus, it was con-
Oral administration of E. longifolia also elicited cluded that E. longifolia may potentially be suit-
sexual arousal in sexually sluggish old male rats able for the management of male infertility.
(Ang et al. 2004b). The results showed that Studies by Chiou and Wu (2012) found that
800 mg/kg of E. longifolia increased yawning by 9-hydroxycanthin-6-one (9-HC-6-one), a
50 % and stretching by 16.7 % in sexually slug- β-carboline alkaloid isolated from E. longifolia,
gish old male rats, by 676–719 % and 31–336 %, could induce penile erection and delayed ejacula-
respectively, in sexually active male rats, and by tion by antagonising the smooth muscle tone of
22–44 % and 75–100 %, respectively, in middle- the rat’s corpus cavernosa as well as seminal ves-
aged, 9-month-old and retired breeders. Zanoli icle probably through interfering with Ca2+
et al. (2009) found that in sexually sluggish rats, mobilisation.
both acute (dosed at 500 and 1000 mg/kg) and Oral administration of a standardised extract
subacute treatments with E. longifolia root pow- F2 (25 mg/kg) of E. longifolia and the aqueous
der significantly reduced ejaculation latencies, extract (250 mg/kg) to male rats significantly
increasing also the percentage of mounting and increased the sperm concentration when com-
ejaculating animals. In addition, the subacute pared with that of the control animals; the effects
administration reduced post-ejaculatory interval. were almost similar in concentration of euryco-
In impotent rats both subacute and subchronic manone, the major and most potent quassinoid
treatments increased the percentage of mounting (Low et al. 2013b). Following treatment with F2,
and ejaculating rats. The effect could be mainly there was a significant increase in the number of
ascribed to increased testosterone levels. The spermatocytes and round spermatids at stage VII
motivational behaviour of sluggish rats during of the spermatogenesis cycle when compared to
the partner preference test was not affected by the that of the control. Plasma levels of testicular
treatments. Studies by Wahab et al. (2010) found testosterone, luteinizing hormone (LH) and
that E. longifolia extract acted as a potential follicle-stimulating hormone (FSH) were higher
264 Simaroubaceae
than those of control plasma, and the oestrogen licular morphological damage. The reversal
level was significantly lower than that of the effect may be attributed to the antiestrogenic
untreated control. The fertility index, fecundity properties of E. longifolia quassinoids.
index and the pup litter size delivered from the Combination treatment of the standardised
females after mating with the males treated with extract (TA164) with artemisinin suppressed P.
F2 were increased. Among the isolated quassi- yoelii infection in the experimental mice. The
noids of F2, eurycomanone and 13α(21)- 4-day suppressive test showed that Eurycoma
dihydroeurycomaone significantly increased the longifolia standardised root extract (TA164) sup-
testosterone level from the Leydig cells of the pressed the parasitemia of Plasmodium yoelii-
testicular interstitial cells cultured in-vitro. The infected mice in a dose-dependent manner (10-,
authors found that eurycomanone dose-depend- 30- and 60-mg/kg BW) by oral and subcutaneous
ently enhanced testosterone steroidogenesis at treatment (Mohd Ridzuan et al. 2007). By oral
the Leydig cells by inhibiting aromatase conver- administration, combination of TA164 at 10-, 30-
sion of testosterone to oestrogen and at a high and 60-mg/kg BW each with artemisinin, respec-
concentration may also involve phosphodiester- tively, showed a significant increase in the
ase inhibition (Low et al. 2013a). The authors parasitemia suppression to 63, 67 and 80 percent
suggested that eurycomanone may be worthy for as compared to artemisinin single treatment
further development as a phytomedicine to treat (31 %). Using subcutaneous administration, at
testosterone-deficient idiopathic male infertility 10-mg/kg BW of TA164 in combination with
and sterility. Treatment of aqueous extract of 1.7-mg/kg BW of artemisinin, gave a suppression
Tongkat Ali root to male rats for 2 weeks of 80 % of infection. Administration of a new
decreased body weight by 5.7 % and omentum herbal combination called Etana, composed of
fat by 31.9 %, and no changes in organ weights five herbal extracts including Panax quinquefo-
were found for the prostate, testes and epididy- lius (ginseng), Eurycoma longifolia (Tongkat
mides (Solomon et al. 2014). Testosterone con- Ali), Epimedium grandiflorum (horny goat
centration increased by 30.2 % and muscle weed), Centella asiatica (gotu kola) and flower
weight also increased, but not significantly. pollen extracts, was found to enhance penile
While sperm concentration, total and progres- erection in male rats (Qinna et al. 2009). The
sive motility and vitality increased significantly, penile erection index (PEI) of Etana was signifi-
mitochondrial membrane potential improved cantly higher than each single component or the
markedly. No detrimental effect could be sum of any two herbal components of Etana.
observed indicating Tongkat Ali to be safe for When compared with sildenafil citrate, Etana
possible treatment of male infertility and ageing induced more pronounced PEI than 0.36 mg/kg,
male problems. but similar to 0.71 mg/kg of sildenafil. Further,
Following intraperitoneal injections at similar full acute and subacute toxicity studies showed
doses of 2.44 μmol/kg/day for 3 consecutive no toxic effects of Etana.
days, eurycomanone displayed comparable
potency with tamoxifen but was more potent than
13α,21-dihydroeurycomanone (quassinoid of E. Antiosteoporotic Activity
longifolia) in the antiestrogenic effect against
17α-ethynylestradiol (EE)-induced uterotrophy Studies showed that both testosterone replace-
of immature rats (Teh et al. 2011a).Treatment ment and E. longifolia supplementation for six
with a standardised quassinoid-rich extract (TAF weeks to orchidectomised aged rats were able to
273) of Eurycoma longifolia root ameliorated restore the bone calcium level, with the former
testosterone-induced reproductive disorders in showing better effects (Shuid et al. 2011). The
female rats (Abdulghani et al. 2012). Upon treat- authors concluded that E. longifolia prevented
ment with TAF 273, fewer animals showed bone calcium loss in orchidectomised rats and
irregular oestrous cycles and there was less fol- therefore had the potential to be used as an
Eurycoma longifolia 265
baumannii and Salmonella typhi) or antifungal when given orally. Its poor oral bioavailability
(Candida albicans, Candida glabrata and may be due to poor membrane permeability in
Candida krusei) effect at concentrations of equal view of its low P value and/or high first-pass
to or less than 50 mg/mL and 10 mg/mL, respec- metabolism.
tively (Tzar et al. 2011) Following intravenous administration of a
standardised extract Fr 2 of Eurycoma longifolia
containing 4.0 % 13α(21)-epoxyeurycomanone
Herb–Drug Interaction Activity (EP), 18.5 % eurycomanone (EN), 0.7 % 13α,21-
dihydroeurycomanone (ED) and 9.5 % of eury-
Salman et al. (2010) found that the bioavailability comanol (EL), EP displayed a relatively longer
of propranolol (a antihypertensive drug) was sig- biological half-life in rat plasma due primarily to
nificantly decreased when consumed together its lower elimination rate constant when com-
with E. longifolia water-based extract in a pared with those of EN (Low et al. 2011).
placebo-controlled randomised single-blind Following oral administration, the absolute bio-
crossover study of 14 healthy non-smoker young availability of EP was 9.5-fold higher than that of
males. The interaction was due to a reduction in EN, not because of chemical degradation since
absorption, rather than an increase in proprano- both quassinoids were stable at the simulated
lol’s metabolism. Although the pharmacodynam- gastric pH of 1. In contrast, EL, being in higher
ics of propranolol was not affected in healthy concentration in the extract than EP, was not
volunteers, caution was still advisable with co- detected in the plasma after oral administration
administration of the drug and the herb. because of substantial degradation by the gastric
Eurycomanone, an active constituent isolated juices after 2 hours. Similarly, ED, being unsta-
from Eurycoma longifolia, was found not to ble at the acidic pH and together with its low con-
inhibit cytochrome P450 (CYP) isoforms centration in Fr 2, was not detectable in the rat
CYP1A2, CYP2A6, CYP2C8, CYP2C9, plasma. Thus, upon oral administration of the
CYP2C19, CYP2E1 and CYP3A4 in in-vitro bioactive standardised extract Fr 2, EP and EN
studies with IC50 values > 250 μg/ml, thus indicat- may be the only quassinoids contributing to the
ing the little likelihood of drug–herb interaction overall antimalarial activity
(Pan et al. 2013).
Toxicity Studies
Pharmacokinetic Studies
An evidence-based systematic review of Tongkat
Studies found that < 1 % of 9-methoxycanthin-6- Ali (Eurycoma longifolia) by the Natural
one, an active compound of Eurycoma longifolia, Standard Research Collaboration consolidated
was found to be absorbed orally (Tan et al. 2002). the safety and efficacy data available in the scien-
A quantification limit of approximately 1.6 ng/ml tific literature (Ulbricht et al. 2013). The topics
of the compound was detected in the rat and covered included written and statistical analysis
human plasma. of clinical trials, plus a compilation of expert
Relatively high plasma eurycomanone con- opinion, folkloric precedent, history, pharmacol-
centrations were detected after an intravenous ogy, kinetics/dynamics, interactions, adverse
injection of 10-mg/kg E. longifolia extract F2 effects, toxicology and dosing.
containing 1.96 mg/kg of the quassinoid (Low Acute toxicity studies of each fraction of the
et al. 2005). However, it declined rapidly to zero 50 % aqueous ethanol extract of Eurycoma longi-
after 8 h. The plasma concentration of the quas- folia root on mice administered orally and brine
sinoid after oral administration was much lower shrimps revealed that the n-butanol fraction was
than after intravenous application in spite of the the most toxic (Chan and Choo 2002).
oral dose being 5 times higher. The results indi- Eurycomanone was identified as the most toxic
cated that eurycomanone was poorly bioavailable component in the n-butanol fraction.
Eurycoma longifolia 269
of the standardised Eurycoma longifolia aqueous folia is that of the Sakai ethnic group in Sumatra
extracts (Physta®) was low and their use posed who use the plant as an amulet to protect people
no genotoxic risks to individuals. from the smallpox virus (Hadiah 1996).
Results of analytical studies showed that 36 % In Peninsular Malaysia and Thailand, the
of the 100 products of Tongkat Ali herbal prepa- roots are used to cure fever, mouth ulcers and
rations possessed 0.52 to 5.30 ppm of mercury intestinal worms (Satayavivad et al. 1998;
and, therefore, did not comply with the quality Jaganath and Ng 2000). It is used in Thai tradi-
requirement for traditional medicines in Malaysia tional medicine as a tonic, aphrodisiac, antican-
(Ang et al. 2004a). Another study showed that cer and antimalarial agent (Kanchanapoom et al.
26 % of Tongkat Ali hitam herbal products pos- 2002). In Vietnam, besides the usual uses (malar-
sessed 0.53–2.35 ppm of mercury and, therefore, ial, dysentery, sexual insufficiency and glandular
did not comply with the quality requirement swelling), a decoction and alcoholic extract of
(≤0.5 ppm) for traditional medicines in Malaysia the roots are used for treatment of rheumatism
(Ang and Lee 2006). (NIMM 2004). In Vietnam, people use the flow-
ers and fruits as a medicine for treating
dysentery.
Traditional Medicinal Uses Fruits of Brucea javanica (L.) Merr.
(‘Ratchadad’ in Thai) and roots of Eurycoma
Eurycoma longifolia is a herbal medicinal plant longifolia Jack (‘Plalaipeag’ in Thai) are used as
of Southeast Asian origin, popularly recognised traditional medicines for the treatment of malar-
as ‘Tongkat Ali’ (Bhat and Karim 2010). Its plant ial fever (Sriwilaijaroen et al. 2010).
parts have been traditionally used for its antima-
larial, aphrodisiac, antidiabetic, antimicrobial
and antipyretic activities, which have also been Other Uses
proven scientifically. ‘Tongkat Ali’ is popularly
taken as a traditional remedy by the ethnic popu- Plant Growth Inhibitory Activity
lation to improve the male libido, sexual prowess
and fertility (Low et al. 2013a). It is traditionally Seven quassinoids from the leaves, including a
used primarily as an aphrodisiac and for improv- new 12-epi-11-dehydroklaineanone, and longi-
ing general health (Ang and Lee 2002a, b, c). lactone, 11-dehydroklaineanone and 15β-O-
Other traditional uses include the treatment of acetyl-14-hydroxyklaineanone showed moderate
aches, persistent fever, malaria, dysentery, growth inhibitory activity of cucumber seedling;
glandular swelling, bleeding, oedema, hyperten- the strongest activity was found in
sion, syphilitic sores and ulcers (Kuo et al. 2004; 14,15β-dihydroxyklaineanone (Jiwajinda et al.
Bedir et al. 2003). Eurycoma longifolia (Tongkat 2001, 2002).
Ali) has been used in the Malay traditional medi-
cine for osteoporotic fracture healing (Abd Jalil
et al. 2012). The inhabitants of some regions of Comments
Sumatra and Kalimantan use the roots an anti-
pyretic (Chan et al. 1995). In Lampung and A digital signal processing (DSP)-based elec-
Belitung, it is used as a medicine for dysentery. tronic taste sensor was developed to authenticate
In Indonesia E. longifolia is used by both large claims of medicinal health food products that
and home industries in the preparation of jamu were enriched with the medicinal herb Eurycoma
(Hadaiah 1996). As is the case with many other longifolia (Abdullah et al. 2004). The system was
species used in the jamu industry, the collection trained to detect E. longifolia into four different
of roots and other plant parts is directly from wild concentrations: 0.01 %, 0.03 %, 0.05 % and
plants. One of the most unique uses for E. longi- 0.08 %. Over 100 samples were tested, from
Eurycoma longifolia 271
which the taste system was able to correctly clas- Selected References
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Medical Glossary
AAD Allergic airway disease, an inflammatory Acetogenins Natural products from the plants
disorder of the airways caused by allergens. of the family Annonaceae, are very potent
AAPH 2,2′-Azobis(2-amidinopropane) dihy- inhibitors of the NADH-ubiquinone reduc-
drochloride, a water-soluble azo compound tase (complex I) activity of mammalian
used extensively as a free radical generator, mitochondria.
often in the study of lipid peroxidation and the Acetyl-CoA carboxylase (ACC) Enzyme that-
characterisation of antioxidants. catalyses the biotin-dependent carboxylation
Abeta aggregation Amyloid beta pro- of acetyl-CoA to produce malonyl-CoA.
tein (Abeta) aggregation is associated with Acetylcholinesterase (AChE) Is an enzyme
Alzheimer’s disease (AD); it is a major com- that degrades (through its hydrolytic activity)
ponent of the extracellular plaque found in the neurotransmitter acetylcholine, producing
AD brains. choline.
Abdominal distension Referring to gener- Acne vulga’ris Also known as chronic acne,
alised distension of most or all of the abdo- usually occurring in adolescence, with com-
men. Also referred to as stomach bloating edones (blackheads), papules (red pimples),
often caused by a sudden increase in fibre nodules (inflamed acne spots) and pustules
from consumption of vegetables, fruits and (small inflamed pus-filled lesions) on the face,
beans. neck and upper part of the trunk.
Ablation therapy The destruction of small Acidosis Increased acidity, an excessively acid
areas of myocardial tissue, usually by appli- condition of the body fluids.
cation of electrical or chemical energy, in the Acquired immunodeficiency syndrome
treatment of some tachyarrhythmias. (AIDS) An epidemic disease caused by
Abortifacient A substance that causes or an infection by human immunodeficiency
induces abortion. virus (HIV-1, HIV-2), retrovirus that causes
Abortivum A substance inducing abortion. immune system failure and debilitation and
Abscess A swollen infected, inflamed area is often accompanied by infections such as
filled with pus in body tissues. tuberculosis.
ABTS 2.2-Azinobis-3-ethylhenthiazoline-6-- Acridone An organic compound based on the
sulphonic acid, a type of mediator in chemical acridine skeleton, with a carbonyl group at the
reaction kinetics of specific enzymes. 9 position.
ACAT Acyl CoA: cholesterol acyltransferase. ACTH Adrenocorticotropic hormone (or cor-
ACE See Angiotensin-converting enzyme. ticotropin), a polypeptide tropic hormone
ACTH (adrenocorticotropic hormone) Also produced and secreted by the anterior pitu-
known as ‘corticotropin’, is a polypeptide itary gland. It palys a role in the synthesis and
tropic hormone produced and secreted by the secretion of gluco- and mineralo-corticoste-
anterior pituitary gland. roids and androgenic steroids.
Activating transcription factor (ATF) A oxygen consumption and coronary blood flow,
protein (gene) that binds to specific DNA while the A2A receptor also has broader anti-
sequences regulating the transfer or transcrip- inflammatory effects throughout the body.
tion of information from DNA to mRNA. These two receptors also have important roles
Activator protein-1 (AP-1) A heterodimeric in the brain, regulating the release of other
protein transcription factor that regulates gene neurotransmitters such as dopamine and glu-
expression in response to a variety of stimuli, tamate, while the A2B and A3 receptors are
including cytokines, growth factors, stress located mainly peripherally and are involved
and bacterial and viral infections. AP-1 in in inflammation and immune responses.
turn regulates a number of cellular processes ADH See Alcohol dehydrogenase.
including differentiation, proliferation and Adipocyte A fat cell involved in the synthesis
apoptosis. and storage of fats.
Actoprotective Increasing the body’s physical Adipocytokine Bioactive cytokines produced
performance. by adipose tissues
Actoprotectors Preparations that increase the Adiponectin A protein in humans that modu-
mental performance and enhance body sta- lates several physiological processes, such
bility against physical loads without increas- as metabolism of glucose and fatty acids and
ing oxygen consumption. Actoprotectors are immune responses.
regarded as a subclass of adaptogens that hold Adipose tissues Body fat, loose connective tis-
a significant capacity to increase physical sue composed of adipocytes (fat cells).
performance. Adoptogen Containing smooth pro-stressors
Acute otitis media (AOM) see Otitis media. which reduce reactivity of host defence sys-
Acyl-CoA dehydrogenases A group of tems and decrease damaging effects of vari-
enzymes that catalyses the initial step in each ous stressors due to increased basal level of
cycle of fatty acid β-oxidation in the mito- mediators involved in the stress response.
chondria of cells. Adrenal glands Star-shaped endocrine glands
Adaptogen A term used by herbalists to refer that sit on top of the kidneys.
to a natural herb product that increases the Adrenalectomised Having had the adrenal
body’s resistance to stresses such as trauma, glands surgically removed.
stress and fatigue. Adrenergic Having to do with adrena-
Adaptogenic Increasing the resistance of the line (epinephrine) and/or noradrenaline
body to stress. (norepinephrine).
Addison’s disease Is a rare endocrine disorder. Adrenergic receptors A class of G protein-
It occurs when the adrenal glands cannot pro- coupled receptors that are targets of the nor-
duce sufficient hormones (corticosteroids). It adrenaline (norepinephrine) and adrenaline
is also known as chronic adrenal insufficiency, (epinephrine).
hypocortisolism or hypocorticism. Adulterant An impure ingredient added into a
Adenocarcinoma A cancer originating in glan- preparation.
dular tissue. Advanced glycation end products
Adenoma A benign tumour from a glandular (AGEs) Resultant products of a chain of
origin. chemical reactions after an initial glycation
Adenoidectomy Surgical removal of the reaction. AGEs may play an important adverse
adenoids. role in process of atherosclerosis, diabetes,
Adenopathy Abnormal enlargement or swell- aging and chronic renal failure.
ing of the lymph node. Aegilops An ulcer or fistula in the inner corner
Adenosine receptors A class of purinergic, of the eye.
G protein-coupled receptors with adenosine Aerophagia Excessive air swallowing.
as endogenous ligand. In humans, there are Afferent Something that so conducts or car-
four adenosine receptors. A1 receptors and A2A ries towards, such as a blood vessel, fibre, or
play roles in the heart, regulating myocardial nerve.
Medical Glossary 279
on subsequent exposures. The term is usually Amastigote Refers to a cell that does not have any
used to refer to hypersensitivity to an environ- flagella, used mainly to describe a certain phase
mental antigen (atopic allergy or contact der- in the life cycle of trypanosome protozoans.
matitis) or to drug allergy. Amenorrhea The condition when a woman
Allodynia A painful response to a normally fails to have menstrual periods.
innocuous stimulus. Amidolytic Cleavage of the amide structure.
Allogeneic Cells or tissues which are geneti- Amoebiasis State of being infected by amoeba
cally different because they are derived from such as Entamoeba histolytica.
separate individuals of the same species. Also Amoebicidal Lethal to amoeba.
refers to a type of immunological reaction AMPK (5′ AMP-activated protein kinase) Or
that occurs when cells are transplanted into a 5’ adenosine monophosphate-activated protein
genetically different recipient. kinase, enzyme that plays a role in cellular
Allografts Or homografts, a graft between energy homeostasis.
individuals of the same species, but of differ- Amygdalitis Inflammation of one or both ton-
ent genotypes. sils, tonsilitis.
Alloknesis Itch produced by innocuous Amyloid beta (Aβ or Abeta) A peptide of
mechanical stimulation. 39–43 amino acids that appear to be the main
Allostasis The process of achieving stability, or constituent of amyloid plaques in the brains of
homeostasis, through physiological or behav- Alzheimer’s disease patients.
ioural change. Amyloidosis A disorder that results from
Alopecia Is the loss of hair on the body. abnormal deposition of the protein, amyloid,
Alopecia areata Type of hair loss that typically in various tissues of the body.
causes patches of baldness. Amyotrophic lateral sclerosis Or ALS, is
ALP See Alkaline phosphatase. a disease of the motor neurons in the brain
Alpha-adrenoceptor Receptors postulated and spinal cord that control voluntary muscle
to exist on nerve cell membranes of the sym- movement.
pathetic nervous system in order to explain Amyotrophy Progressive wasting of muscle
the specificity of certain agents that affect tissues. adj. amyotrophic.
only some sympathetic activities (such as Anaemia A blood disorder in which the blood is
vasoconstriction and relaxation of intes- deficient in red blood cells and in haemoglobin.
tinal muscles and contraction of smooth Anaesthesia Condition of having sansation
muscles). temporarily suppressed.
Alpha amylase (α-amylase) A major form of Anaesthetic A substance that decreases par-
amylase found in humans and other mammals tially or totally nerve the sense of pain.
that cleaves alpha-bonds of large, alpha-linked Analeptic A central nervous system (CNS)
polysaccharides, such as starch and glycogen, stimulant medication.
yielding glucose and maltose. Analgesia Term describing relief, reduction or
ALT See Alanine transaminase. suppression of pain. adj. analgetic.
Alterative A medication or treatment which Analgesic A substance that relieves or reduces
gradually induces a change and restores healthy pain.
functions without sensible evacuations. Anamnesis Patient’s account of their medical
Alveolar macrophage A vigorously phago- history.
cytic macrophage on the epithelial surface Anaphoretic An antiperspirant.
of lung alveoli that ingests carbon and other Anaphodisiac Or antiaphrodisiac is something
inhaled particulate matters. Also called conio- that reduces or blunts the libido.
phage or dust cell. Anaphylaxis A severe, life-threatening allergic
Alzheimer’s disease A degenerative, organic, response that may be characterised by symp-
mental disease characterised by progressive toms such as reduced blood pressure, wheez-
brain deterioration and dementia, usually ing, vomiting or diarrhoea.
occurring after the age of 50. Anaphylactic adj. see Anaphylaxis.
Medical Glossary 281
Anaphylotoxins Are fragments (C3a, C4a or (RAS) which mediates extracellular volume
C5a) that are produced during the pathways (i.e. that of the blood plasma, lymph and inter-
of the complement system. They can trigger stitial fluid) and arterial vasoconstriction.
release of substances of endothelial cells, mast Anglioplasty Medical procedure used to open
cells or phagocytes, which produce a local obstructed or narrowed blood vessel resulting
inflammatory response. usually from atherosclerosis.
Anaplasia A reversion of differentiation in Anguillulosis A parasitosis caused by the
cells and is characteristic of malignant neo- intestinal nematode Strongyloides stercoralis
plasms (tumours). (round worm).
Anaplastic adj. see Anaplasia. Anisakiasis A human parasitic infection of the
Anasarca Accumulation of great quantity of gastrointestinal tract caused by the consump-
fluid in body tissues. tion of raw or undercooked seafood contain-
Anencephaly A cephalic disorder that results ing larvae of the nematode Anisakis simplex.
from a neural tube defect that occurs when the Anisonucleosis A morphological manifesta-
cephalic (head) end of the neural tube fails to tion of nuclear injury characterised by varia-
close, resulting in the absence of a major por- tion in the size of the cell nuclei.
tion of the brain, skull and scalp. Ankylosing spondylitis (AS) Is a type of
Androgen Male sex hormone in vertebrates. inflammatory arthritis that targets the joints of
Androgens may be used in patients with breast the spine.
cancer to treat recurrence of the disease. Annexin V or annexin A5 Is a member of the
Androgenic alopecia Hair loss in men and annexin family of intracellular proteins that
women. binds to phosphatidylserine (PS) in a calcium-
Android adiposity Centric fat distribution pat- dependent manner.
terns with increased disposition towards the Annexitis Also called adnexitis, a pelvic
abdominal area, visceral fat—apple shaped. inflammatory disease involving the inflamma-
cf. gynoid adiposity. tion of the ovaries or fallopian tubes.
Andrology Branch of medicine concerned Anodyne A substance that relieves or soothes
with the reproductive diseases in men. pain by lessening the sensitivity of the brain
Aneugen An agent that affects cell division or nervous system. Also called an analgesic.
and the mitotic spindle apparatus, causing the Anoikis Apoptosis that is induced by inade-
loss or gain of whole chromosomes, thereby quate or inappropriate cell-matrix interactions.
inducing aneuploidy. adj. aneugenic. Anophthalmia Medical term for the absence
Angina pectoris, angina Chest pain or chest of one or both eyes.
discomfort that occurs when the heart muscle Anorectal Relating to the rectum and anus.
does not get enough blood. Anorectics Appetite suppressants, substances
Angioedema Rapid swelling of the dermis, which reduce the desire to eat. Used on a
subcutaneous tissues, mucosa and submucosal short-term basis clinically to treat obesity.
tissues caused by small blood vessels leaking Also called anorexigenics.
fluid into these tissues. Anorexia Lack or loss of desire to eat.
Angiogenic adj. see Angiogenesis. Anorexia nervosa Is a psychiatric disorder
Angiogenesis A physiological process involv- characterised by an unrealistic fear of weight
ing the growth of new blood vessels from pre- gain, self-starvation and conspicuous distor-
existing vessels. tion of body image.
Angiotensin An oligopeptide hormone in the Anorexic Having no appetite to eat.
blood that causes blood vessels to constrict Anorexigenics See Anorectics.
and drives blood pressure up. It is part of the Anosmia Inabity to perceive odour, reduced
renin–angiotensin system. sense of smell.
Angiotensin-converting enzyme (ACE) An Anoxia Absence of oxygen supply.
exopeptidase, a circulating enzyme that partic- Antagonist A substance that acts against and
ipates in the body’s renin–angiotensin system blocks an action.
282 Medical Glossary
Antalgic A substance used to relive a painful ity to inhibit the growth of or to kill other
condition. microorganisms.
Antecubital vein This vein is located in the Antiblennorrhagic A substance that treats
antecubital fossa—the area of the arm in front blenorrhagia a conjunctival inflammation
of the elbow. resulting in mucus discharge.
Anterior uveitis Is the most common form of Antibody A gamma globulin protein produced
ocular inflammation that often causes a pain- by a kind of white blood cell called the plasma
ful red eye. cell in the blood used by the immune sys-
Anthelmintic An agent or substance that is tem to identify and neutralise foreign objects
destructive to worms and used for expulsion (antigen).
of internal parasitic worms in animals and Anticarcinomic A substance that kills or
humans. inhibits carcinomas (any cancer that arises in
Anthocyanins A subgroup of antioxidant fla- epithelium/tissue cells).
vonoids, are glucosides of anthocyanidins, Anticephalalgic Headache relieving or
which are beneficial to health. They occur preventing.
as water-soluble vacuolar pigments that may Anticestodal A chemical destructive to
appear red, purple or blue according to pH in tapeworms.
plants. Anticholesterolemic A substance that can pre-
Anthrax A bacterial disease of cattle and ship vent the build-up of cholesterol.
that can be transmitted to man though unpro- Anticlastogenic Having a suppressing effect of
cessed wool. chromosomal aberrations.
Anthropometric Pertaining to the study of Anticoagulant A substance that thins the blood
human body measurements. and acts to inhibit blood platelets from stick-
Antiamoebic A substance that destroys or sup- ing together.
presses parasitic amoebae. Antidepressant A substance that suppresses
Antiamyloidogenic Compounds that inhibit depression or sadness.
the formation of Alzheimer’s β-amyloid fibrils Antidiabetic A substance that prevents or alle-
(fAβ) from amyloid β-peptide (Aβ) and desta- viates diabetes. Also called antidiabetogenic.
bilise fAβ. Antidiarrhoeal Having the property of stop-
Antianaphylactic Agent that can prevent the ping or correcting diarrhoea, an agent having
occurrence of anaphylaxis (life-threatening such action.
allergic response). Antidipsotropic Antialcohol abuse; medica-
Antiangiogenic A drug or substance used to tion to reduce alcohol consumption.
stop the growth of tumours and progression of Antidote A remedy for counteracting a poison.
cancers by limiting the pathologic formation Antidopaminergic A term for a chemical
of new blood vessels (angiogenesis). that prevents or counteracts the effects of
Antiarrhythmic A substance to correct irregu- dopamine.
lar heartbeats and restore the normal rhythm. Antidrepanocytary Anti-sickle cell anaemia.
Antiasmathic Drug that treats or ameliorates Antidysenteric An agent used to reduce or
asthma. treat dysentery and diarrhoea.
Antiatherogenic That protects against athero- Antidyslipidemic Agent that will reduce the
genesis, the formation of atheromas (plaques) abnormal amount of lipids and lipoproteins in
in arteries. the blood.
Antibacterial Substance that kills or inhibits Anti-oedematous Reduces or suppresses
bacteria. oedema.
Antibilious An agent or substance which helps Antiemetic An agent that stops vomiting and
remove excess bile from the body. nausea.
Antibiotic A chemical substance produced Anti-epileptic A drug used to treat or prevent
by a microorganism which has the capac- convulsions, anticonvulsant.
Medical Glossary 283
Antioestrogen A substance that inhibits the sense RNA sequence, translation of the
biological effects of female sex hormones. mRNA into the protein is blocked. This may
Antiophidian Antivenoms of snake. slow or halt the growth of cancer cells.
Antiosteoporotic Substance that can prevent Antiseptic Preventing decay or putrefaction, a
osteoporosis. substance inhibiting the growth and develop-
Antiovulatory Substance suppressing ment of microorganisms.
ovulation. Anti-sickling agent An agent used to pre-
Antioxidant A chemical compound or sub- vent or reverse the pathological events lead-
stance that inhibits oxidation and protects ing to sickling of erythrocytes in sickle cell
against free radical activity and lipid oxidation conditions.
such as vitamin E, vitamin C or beta-carotene Antispasmodic A substance that relieves
(converted to vitamin B) and carotenoids and spasms or inhibits the contraction of smooth
flavonoids which are thought to protect body muscles; smooth muscle relaxant, muscle
cells from the damaging effects of oxidation. relaxer.
Many foods including fruit and vegetables Antispermatogenic Preventing or suppressing
contain compounds with antioxidant proper- the production of semen or spermatozoa.
ties. Antioxidants may also reduce the risks of Antisudorific See Antiperspirant.
cancer and age-related macular degeneration Antisyphilitic A drug (or other chemical
(AMD). agents) that is effective against syphilis.
Antipaludic Antimalarial. Antithermic A substance that reduces fever
Antiperiodic Substance that prevents the recur- and temperature. Also known as antipyretic.
rence of symptoms of a disease, e.g. malaria. Antithrombotic Preventing or interfering with
Antiperspirant A substance that inhibits sweat- the formation of thrombi.
ing. Also called antisudorific, anaphoretic. Antitoxin An antibody with the ability to neu-
Antiphlogistic A traditional term for a tralise a specific toxin.
substance used against inflammation, an Antitumoral Substance that acts against the
anti-inflammatory. growth, development or spread of a tumour.
Antiplatelet agent Drug that decreases platelet Antitussive A substance that depresses
aggregation and inhibits thrombus formation. coughing.
Antiplasmodial Suppressing or destroying Antiulcerogenic An agent used to protect
plasmodia. against the formation of ulcers or is used for
Antiproliferative Preventing or inhibiting the the treatment of ulcers.
reproduction of similar cells. Antivenin An agent used against the venom of
Antiprostatic Drug to treat the prostate. a snake, spider or other venomous animals or
Antiprotozoal Suppressing the growth or insects.
reproduction of protozoa. Antivinous An agent or substance that treats
Antipruritic Alleviating or preventing itching. addiction to alcohol.
Antipyretic A substance that reduces fever or Antiviral Substance that destroys or inhibits
quells it. Also known as antithermic. the growth and viability of infectious viruses.
Antirheumatic Relieving or preventing Antivomitive A substance that reduces or sup-
rheumatism. presses vomiting.
Antiscorbutic A substance or plant rich in Antizygotic See Antifertility.
vitamin C that is used to counteract scurvy. Anuria Absence of urine production and
Antisecretory Inhibiting or diminishing excretion. adj. anuric.
secretion. Anxiogenic Substance that causes anxiety.
Antisense Refers to antisense RNA strand Anxiolytic A drug prescribed for the treatment
because its sequence of nucleotides is the of symptoms of anxiety.
complement of message sense. When mRNA APAF-1 Apoptotic protease-activating factor 1.
forms a duplex with a complementary anti-
Medical Glossary 285
Apelin Also known as APLN, a peptide which aPPT (activated partial thromboplastin
in humans is encoded by the APLN gene. time) A blood test, a measure of the part of
Aperient A substance that acts as a mild laxa- the blood clotting pathway.
tive by increasing fluids in the bowel. Apurinic lyase A DNA enzyme that catalyses
Aperitif An appetite stimulant. a chemical reaction.
Aphonia Loss of the voice resulting from dis- Arachidonate cascade Includes the cyclooxy-
ease, injury to the vocal cords, or various psy- genase (COX) pathway to form prostanoids
chological causes, such as hysteria. and the lipoxygenase (LOX) pathway to gen-
Aphrodisiac An agent that increases sexual erate several oxygenated fatty acids, collec-
activity and libido and/or improves sexual tively called eicosanoids.
performance. ARE Antioxidant response element, is a tran-
Aphthae White, painful oral ulcer of unknown scriptional control element that mediates
cause. expression of a set of antioxidant proteins.
Aphthous ulcer Also known as a canker sore, Ariboflavinosis A condition caused by the
is a type of oral ulcer, which presents as a dietary deficiency of riboflavin that is char-
painful open sore inside the mouth or upper acterised by mouth lesions, seborrhea and
throat. vascularisation.
Aphthous stomatitis A canker sore, a type of Aromatase An enzyme involved in the produc-
painful oral ulcer or sore inside the mouth or tion of oestrogen that acts by catalysing the
upper throat, caused by a break in the mucous conversion of testosterone (an androgen) to
membrane. Also called aphthous ulcer. estradiol (an oestrogen). Aromatase is located
Aplastic anaemia (AA) is a disease in which in oestrogen-producing cells in the adrenal
the bone marrow and the blood stem cells that glands, ovaries, placenta, testicles, adipose
reside there are damaged and do not make (fat) tissue and brain.
enough new blood cells. Aromatic Having a pleasant, fragrant odour.
Apnoea Suspension of external breathing. Aromatherapy A form of alternative medicine
Apoprotein The protein moiety of a molecule that uses volatile liquid plant materials, such
or complex, as of a lipoprotein. as essential oils and other scented compounds
Apolipoprotein A-I (APOA1) A major protein from plants for the purpose of affecting a per-
component of high-density lipoprotein (HDL) son’s mood or health.
in plasma. The protein promotes cholesterol ARPE-19 cells A human retinal pigment epi-
efflux from tissues to the liver for excretion. thelial cell line with differentiated properties.
Apolipoprotein B (APOB) Is the primary apo- Arrhythmias Abnormal heart rhythms that can
lipoprotein of low-density lipoproteins (LDL cause the heart to pump less effectively. Also
or ‘bad cholesterol’), which is responsible for called dysrhythmias.
carrying cholesterol to tissues. Arsenicosis See Arsenism.
Apolipoprotein E (APOE) The apolipoprotein Arsenism An incommunicable disease result-
found on intemediate density lipoprotein and ing from the ingestion of groundwater con-
chylomicron that binds to a specific receptor taining unsafe levels of arsenic, also known as
on liver and peripheral cells. arsenicosis.
Apoplexy Unconsciousness or incapacity of Arteriogenic erectile dysfunction A penis
the brain to function resulting from a cerebral dysfunction caused by the narrowing of the
haemorrhage or stroke. arteries in the penis, decreasing blood inflow
Apoptogenic Ability to cause death of cells. to it, thus making erection impossible.
Apoptosis Death of cells. Arteriosclerosis Imprecise term for various
Appendicitis Is a condition characterised by disorders of arteries, particularly hardening
inflammation of the appendix. Also called due to fibrosis or calcium deposition, often
epityphlitis. used as a synonym for atherosclerosis.
Appetite stimulant A substance to increase or Arthralgia Is pain in the joints from many pos-
stimulate the appetite. Also called aperitif. sible causes.
286 Medical Glossary
Arthritis Inflammation of the joints of the with excessive amounts of mucus, often in
body. response to one or more triggers.
Arthrodynia An affection characterised by Astringent A substance that contracts blood
pain in or about a joint. vessels and certain body tissues (such as
Arthus reaction An allergic reaction of the mucous membranes) with the effect of reduc-
immediate hypersensitive type that results ing secretion and excretion of fluids and/or
from the union of antigen and antibody, with has a drying effect.
complement present in blood vessel walls. Astrocytes Collectively called astroglia, are
Aryl hydrocarbon receptor (AhR) A characteristic star-shaped glial cells in the
ligand-activated transcription factor best brain and spinal cord.
known for mediating the toxicity of dioxin Ataxia (Loss of coordination) results from the
and other exogenous contaminants and is degeneration of nerve tissue in the spinal cord
responsible for their toxic effects, including and of nerves that control muscle movement
immunosuppression. in the arms and legs.
ASATor AST Aspartate aminotransferase; see Ataxic polyneuropathy Is a syndrome char-
Aspartate transaminase. acterised by problems with coordination and
ASBT Apical sodium-dependent bile acid balance (sensory ataxia) and disturbances in
transporter, belongs to the solute carrier fam- nerve function (sensory neuropathy), bilat-
ily (SLC) of transporters and is an important eral optic atrophy and bilateral sensorineural
carrier protein expressed in the small intestine. deafness.
Ascaris A genus of parasitic intestinal round Ataxia telangiectasia and Rad3-related pro-
worms. tein (ATR) Also known as serine/threonine-
Ascites Abnormal accumulation of fluid within protein kinase ATR, FRAP-related protein 1
the abdominal or peritoneal cavity. (FRP1), is an enzyme encoded by the ATR
Ascorbic acid See Vitamin C. gene. It is involved in sensing DNA damage
Aspartate transaminase (AST) Also called and activating the DNA damage checkpoint,
serum glutamic oxaloacetic transaminase leading to cell cycle arrest
(SGOT) or aspartate aminotransferase (ASAT) Atelectasis The collapse or closure of the
is similar to ALT in that it is another enzyme lung resulting in reduced or absent gas
associated with liver parenchymal cells. It is exchange.
increased in acute liver damage but is also pres- ATF-2 Activating transcription factor 2.
ent in red blood cells and cardiac and skeletal Athlete’s foot A contagious skin disease
muscle and is therefore not specific to the liver. caused by parasitic fungi affecting the foot
Asphyxia Failure or suppression of the respi- and hands, causing itching, blisters and crack-
ratory process due to obstruction of airflow ing. Also called dermatophytosis.
to the lungs or due to the lack of oxygen in Atherogenic Having the capacity to start or
inspired air. accelerate the process of atherogenesis.
Asphyxiation The process of undergoing Atherogenesis The formation of lipid deposits
asphyxia. in the arteries.
Asthenia A nonspecific symptom character- Atheroma A deposit or degenerative accumu-
ised by loss of energy, strength and feeling of lation of lipid-containing plaques on the inner-
weakness. most layer of the wall of an artery.
Asthenopia Weakness or fatigue of the eyes, Atherosclerosis The condition in which an
usually accompanied by headache and dim- artery wall thickens as the result of a build-up
ming of vision. adj. asthenopic. of fatty materials such as cholesterol.
Asthenozoospermia (asthenosper- Atherothrombosis Medical condition charac-
mia) Reduced sperm motility. terised by an unpredictable, sudden disruption
Asthma A chronic illness involving the respi- (rupture or erosion/fissure) of an atheroscle-
ratory system in which the airway occasion- rotic plaque, which leads to platelet activation
ally constricts, becomes inflamed and is lined and thrombus formation.
Medical Glossary 287
Athymic mice Laboratory mice lacking a thy- Autolysin An enzyme that hydrolyses and
mus gland. destroys the components of a biological cell
Atonic Lacking normal tone or strength. or a tissue in which it is produced.
Atony Insufficient muscular tone. Autonomic disorder A neurological disease in
Atopic dermatitis An inflammatory, non- which the autonomic nervous system ceases to
contagious, pruritic skin disorder of unknown function properly.
aetiology; often called eczema. Autonomic neuropathy Is a nerve disorder
Atresia A congenital medical condition in due to damage to the autonomic nerves that
which a body orifice or passage in the body is affects involuntary body functions, including
abnormally closed or absent. heart rate, blood pressure, perspiration and
Atretic follicle Follicular atresia is the break- digestion.
down of the ovarian follicles. Autophagy Digestion of the cell contents by
Atretic ovarian follicles An involuted or enzymes in the same cell.
closed ovarian follicle. Autopsy Examination of a cadaver to deter-
Atrial fibrillation Is the most common car- mine or confirm the cause of death.
diac arrhythmia (abnormal heart rhythm) and Avenanthramides Low molecular weight, sol-
involves the two upper chambers (atria) of the uble phenolic compounds found in oats.
heart; the most serious consequence of atrial Avidity index Describes the collective inter-
fribillation is ischemic stroke. actions between antibodies and a multivalent
Atrioventricular node A node of specialised antigen.
heart muscle located in the septal wall of the Avulsed teeth Is tooth that has been knocked
right atrium; receives impulses from the sino- out.
atrial node and directs them to the walls of the Ayurvedic Traditional Hindu system of
ventricles. medicine based largely on homeopathy and
Attention-deficit hyperactivity disorder naturopathy.
(ADHD, ADD or AD/HD) Is a neurobe- Azoospermia Is the medical condition of a
havioural developmental disorder, primarily male not having any measurable level of
characterised by the coexistence of attentional sperm in his semen.
problems and hyperactivity. Azotaemia A higher than normal blood level
Auditory brainstem response (ABR) Also of urea or other nitrogen-containing com-
called brainstem evoked response (BSER), is pounds in the blood.
an electrical signal evoked from the brainstem B Cell-activating factor (BAFF) Also called
of a human by the presentation of a sound tumour necrosis factor ligand superfamily
such as a click. member 13B. It plays an important role in the
Augmerosen A drug that may kill cancer cells proliferation and differentiation of B cells.
by blocking the production of a protein that Babesia A protozoan parasite (malaria-like)
makes cancer cells live longer. Also called of the blood that causes a haemolytic disease
bcl-2 antisense oligonucleotide. known as babesiosis.
Auricular Of or relating to the auricle or the Babesiosis Malaria-like parasitic disease
ear in general. caused by Babesia, a genus of protozoal
Aurones [2-Benzylidenebenzofuran-3(2H)- piroplasms.
ones] are the secondary plant metabo- Back tonus Normal state of balanced tension
lites and are a subgroup of flavonoids. See in the tissues of the back.
Flavonoids. Bactericidal Lethal to bacteria.
Autoantibodies Antibodies manufactured by BAFF A cytokine that belongs to the tumour
the immune system that mistakenly target necrosis factor (TNF) ligand family.
and damage specific tissues and organs of the Balanitis Is an inflammation of the glans
body. (head) of the penis.
288 Medical Glossary
BALB/c mice Balb/c mouse was developed in differentiation and apoptosis of some tumour
1923 by McDowell. It is a popular strain and cell types, particularly in early stages of
is used in many different research disciplines, tumorigenesis, and enhances immune system
but most often in the production of monoclo- activity by stimulating the release of natural
nal antibodies. killer cells, lymphocytes and monocytes.
Balm Aromatic oily resin from certain trees Beta-catenin Is a multifunctional oncogenic
and shrubs used in medicine. protein that contributes fundamentally to cell
Barbiturates Are drugs that act as central development and biology. It has been impli-
nervous system depressants and can there- cated as an integral component in the Wnt sig-
fore produce a wide spectrum of effects, nalling pathway.
from mild sedation to total anaesthesia. They Beta cells A type of cell in the pancreas in
are also effective anxiolytics, hypnotics and areas called the islets of Langerhans.
anticonvulsants. Beta glucans Polysaccharides of D-glucose
Baroreceptor A type of interoceptor that is monomers linked by β-glycosidic bonds,
stimulated by pressure changes, as those in (1→3), (1→4)-β-D-glucan, soluble, viscous
blood vessel wall. component of fibres found in cereals like oats.
Barrett’s oesophagus (Barrett oesophagi- Beta-thalassemia An inherited blood disorder
tis) A disorder in which the lining of the that reduces the production of haemoglobin.
oesophagus is damaged by stomach acid. Beta-lactamase Enzymes produced by some
Basophil A type of white blood cell with coarse bacteria that are responsible for their resistance
granules within the cytoplasm and a bilobate to beta-lactam antibiotics like penicillins.
(two-lobed) nucleus. BHT Butylated hydroxytoluene (phenolic
Bax/Bad Proapoptotic proteins. compound), an antioxidant used in foods,
BCL-2 A family of apoptosis regulator pro- cosmetics, pharmaceuticals and petroleum
teins in humans encoded by the B cell lym- products.
phoma 2 (BCL-2) gene. BID The only known Bcl-2 family member
BCL-2 antisense oligonucleotide See that can function as an agonist of proapoptotic
Augmereson. Bcl-2-related proteins such as Bax and Bak.
BCR/ABL A chimeric oncogene, from fusion Bifidobacterium Is a genus of Gram-positive,
of BCR and ABL cancer genes associated nonmotile, often branched anaerobic bacte-
with chronic myelogenous leukaemia. ria. Bifidobacteria are one of the major gen-
Bechic A remedy or treatment of cough. era of bacteria that make up the gut flora.
Bed nucleus of the stria terminalis Bifidobacteria aid in digestion are associated
(BNST) Act as a relay site within the hypo- with a lower incidence of allergies and also
thalamic-pituitary-adrenal axis and regulate prevent some forms of tumour growth. Some
its activity in response to acute stress. bifidobacteria are being used as probiotics.
Belching, or burping Refers to the noisy Bifidogenic Promoting the growth of (benefi-
release of air or gas from the stomach through cial) bifidobacteria in the intestinal tract.
the mouth. Bile Fluid secreted by the liver and discharged
Beriberi Is a disease caused by a deficiency of into the duodenum where it is integral in the
thiamine (vitamin B1) that affects many sys- digestion and absorption of fats.
tems of the body, including the muscles, heart, Bilharzia, bilharziosis See Schistosomiasis.
nerves and digestive system. Biliary Relating to the bile or the organs in
Beta-carotene Naturally occurring retinol which the bile is contained or transported.
(vitamin A) precursor obtained from certain Biliary infections Infection of organ(s) associ-
fruits and vegetables with potential antineo- ated with bile, comprise:
plastic and chemopreventive activities. As an (a) acute cholecystitis, an acute inflammation of
antioxidant, beta-carotene inhibits free radical the gallbladder wall
damage to DNA. This agent also induces cell (b) cholangitis, inflammation of the bile ducts
Medical Glossary 289
Biliousness Old term used in the eighteenth Blood stasis syndrome Blood stagnation or
and nineteenth centuries pertaining to bad slowing of blood, an important underlying
digestion, stomach pains, constipation and pathology of many disease processes accord-
excessive flatulence. ing to traditional Chinese medicine.
Bilirubin A breakdown product of haem (a BMPs (bone morphogenetic proteins) A
part of haemoglobin in red blood cells) pro- family of secreted signalling molecules that
duced by the liver that is excreted in bile which can induce ectopic bone growth.
causes a yellow discoloration of the skin and BNIP3 A pro-apoptotic BH3-only protein
eyes when it accumulates in those organs. which is associated with mitochondrial dys-
Biotin Also known as vitamin B7. See Vitamin function and cell death.
B7. Boil Localised pyrogenic, painful infection,
Bitter A medicinal agent with a bitter taste and originating in a hair follicle.
used as a tonic, alterative or appetiser. Borborygmus Rumbling noise caused by the
Blackhead See Comedone. muscular contractions of peristalsis, the pro-
Blackwater fever Dangerous complication of cess that moves the contents of the stomach
malarial whereby the red blood cells burst in and intestines downwards.
the blood stream (haemolysis) releasing hae- Bowman–Birk inhibitors Type of serine pro-
moglobin directly into the blood. teinase inhibitor.
Blain See Chilblain. Bouillon A broth in French cuisine.
Blastocyst Blastocyst is an embryonic struc- Bradicardia As applied to adult medicine,
ture formed in the early embryogenesis of is defined as a resting heart rate of under 60
mammals, after the formation of the morula, beats per minute.
but before implantation. Bradyphrenia Referring to the slowness of
Blastocystotoxic Agent that suppresses further thought common to many disorders of the
development of the blastocyst through to the brain.
ovum stage. Brain-derived neutrophic factor (BDNF) A
Blebbing Bulging, e.g. membrane blebbing protein member of the neutrophin family that
also called membrane bulging or ballooning. plays an important role in the growth, mainte-
Bleeding diathesis Is an unusual susceptibility nance, function and survival of neurons. The
to bleeding (haemorrhage) due to a defect in protein molecule is involved in the modulation
the system of coagulation. of cognitive and emotional functions and in
Blennorrhagia Gonorrhoea. the treatment of a variety of mental disorders.
Blennorrhoea Inordinate discharge of mucus, Bright’s disease Chronic nephritis.
especially a gonorrhoeal discharge from the Bronchial inflammation See Bronchitis.
urethra or vagina. Bronchiectasis A condition in which the
Blepharitis Inflammation of the eyelids. airways within the lungs (bronchial tubes)
Blepharospasm Involuntary twitching, blink- become damaged and widened.
ing closure or squeezing of the eyelids. Bronchitis Is an inflammation of the main air
Blister Thin vesicle on the skin containing passages (bronchi) to the lungs.
serum and caused by rubbing, friction or Bronchoalveolar lavage (BAL) A medical
burn. procedure in which a bronchoscope is passed
Blood–brain barrier (BBB) Is a separation through the mouth or nose into the lungs and
of circulating blood and cerebrospinal fluid fluid is squirted into a small part of the lung
(CSF) in the central nervous system (CNS). and then recollected for examination.
It allows essential metabolites, such as oxy- Bronchopneumonia Or bronchial pneumonia;
gen and glucose, to pass from the blood to the inflammation of the lungs beginning in the ter-
brain and central nervous system (CNS) but minal bronchioles.
blocks most molecules that are more massive Broncho-pulmonary Relating to the bronchi
than about 500 Da. and lungs.
290 Medical Glossary
bolic reactions that involve the oxidation of cGMP Cyclic guanosine monophosphate is
one molecule and the reduction of another. a cyclic nucleotide derived from guanosine
Cellulitis A bacterial infection of the skin that triphosphate (GTP). cGMP is a common reg-
tends to occur in areas that have been dam- ulator of ion channel conductance, glycoge-
aged or inflamed. nolysis and cellular apoptosis. It also relaxes
Central nervous system Part of the vertebrate smooth muscle tissues.
nervous system comprising the brain and spi- CGRP calcitonin gene-related peptide A
nal cord. vasodilator neuropeptide that is expressed in a
Central serous chorioretinopathy (CSCR) subgroup of small neurons in the dorsal root,
is a disease in which a serous detachment of trigeminal and vagal ganglia. This neuropep-
the neurosensory retina occurs over an area of tide has been postulated to play a role in the
leakage from the choriocapillaris through the pathophysiology of migraine.
retinal pigment epithelium. Chalcones A subgroup of flavonoids.
Central venous catheter A catheter placed Chancre A painless lesion formed during the
into the large vein in the neck, chest or groin. primary stage of syphilis.
Cephalagia Pain in the head, a headache. Chaperones Are proteins that assist the non-
Cephalic Relating to the head. covalent folding or unfolding and the assem-
Ceramide oligosides Oligosides with an bly or disassembly of other macromolecular
N-acetyl-sphingosine moiety. structures.
Cercariae A free swimming larva of the para- Chemoembolisation A procedure in which
sitic schistosome worm that has a tail and the blood supply to the tumour is blocked
suckers on its head for penetration into a host. surgically or mechanically and antican-
Cerebral embolism A blockage of blood flow cer drugs are administered directly into the
through a vessel in the brain by a blood clot tumour.
that formed elsewhere in the body and trav- Chemokines Are chemotactic cytokines,
elled to the brain. which stimulate migration of inflammatory
Cerebral ischemia Is the localised reduc- cells towards tissue sites of inflammation.
tion of blood flow to the brain or parts of the Chemonociceptors Nociceptors or sensory
brain due to arterial obstruction or systematic pheripheral neurons that are sensitive to
hyperfusion. chemical stimuli.
Cerebral infarction Is the ischemic kind of Chemosensitiser A drug that makes tumour
stroke due to a disturbance in the blood ves- cells more sensitive to the effects of
sels supplying blood to the brain. chemotherapy.
Cerebral tonic Substance that can alleviate Chemosis Oedema of the conjunctiva of the
poor concentration and memory, restlessness, eye.
uneasiness and insomnia. Chickenpox Is also known as varicella and
Cerebrosides Are glycosphingolipids which is a highly contagious illness caused by pri-
are important components in animal muscle mary infection with varicella zoster virus
and nerve cell membranes. (VZV). The virus causes red, itchy bumps on
Cerebrovascular disease Is a group of brain the body.
dysfunctions related to disease of the blood Chilblains Small, itchy, painful lumps that
vessels supplying the brain. develop on the skin. They develop as an
Cervical spondylotic myelopathy A com- abnormal response to cold. Also called per-
mon cause of spinal cord dysfunction in older niosis or blain.
persons. Chlorosis Iron deficiency anaemia character-
Cerumen Ear wax, a yellowish waxy sub- ised by greenish yellow colour.
stance secreted in the ear canal of humans and Cholagogue Is a medicinal agent which pro-
other mammals. motes the discharge of bile from the system.
cFLIP Cellular FLICE-inhibitory protein, an Cholecalcifereol A form of vitamin D, also
inhibitor of death ligand-induced apoptosis. called vitamin D3. See Vitamin D.
294 Medical Glossary
Clastogen Is an agent that can cause one of Coma A state of unconsciousness from which
two types of structural changes and breaks in a patient cannot be aroused.
chromosomes that results in the gain, loss, or Comedone A blocked, open sebaceous gland
rearrangements of chromosomal segments. where the secretions oxidise, turning black.
adj. clastogenic. Also called blackhead.
Claudication Limping, impairment in walking. Comitogen Agent that is considered not to
Climacterium Refers to menopause and the induce cell growth alone but to promote the
bodily and mental changes associated with it. effect of the mitogen.
Clonic seizures Consist of rhythmic jerking Concoction A combination of crude ingredi-
movements of the arms and legs, sometimes ents that is prepared or cooked together.
on both sides of the body. Condyloma, condylomata acuminata Genital
Clonus A series of involuntary muscular con- wart, venereal wart, anal wart or anogenital
tractions and relaxations. wart, a highly contagious sexually transmitted
Clyster Enema. infection caused by epidermotropic human
C-myc Codes for a protein that binds to the papillomavirus (HPV).
DNA of other genes and is therefore a tran- Congestive heart failure Heart failure in
scription factor. which the heart is unable to maintain adequate
CNS depressant Anything that depresses, or circulation of blood in the tissues of the body
slows, the sympathetic impulses of the cen- or to pump out the venous blood returned to it
tral nervous system (i.e. respiratory rate, heart by the venous circulation.
rate). Conglutination Becoming stuck together.
Coagulopathy A defect in the body’s mecha- Conjunctival hyperemia Enlarged blood ves-
nism for blood clotting, causing susceptibility sels in the eyes.
to bleeding. Conjunctivitis Sore, red and sticky eyes
Cobalamin Vitamin B12. See Vitamin B12. caused by eye infection.
Co-carcinogen A chemical that promotes the Conn’s syndrome Extremely rare condi-
effects of a carcinogen in the production of tion characterised by adenoma, carcinoma or
cancer. hyperplasia of the zona glomerulosa of the
Cold An acute inflammation of the mucous adrenal cortex, resulting in excessive produc-
membrane of the respiratory tract, especially tion of aldosterone and leading to sodium
of the nose and throat caused by a virus and retention and hydrogen loss.
accompanied by sneezing and coughing. Constipation A very common gastrointestinal
Collagen Protein that is the major constituent disorder characterised by the passing of hard,
of cartilage and other connective tissues; com- dry bowel motions (stools) and difficulty of
prises the amino acids, hydroxyproline, pro- bowel motion.
line, glycine and hydroxylysine. Constitutive androstane receptor (CAR,
Collagenases Enzymes that break the peptide NR113) Is a nuclear receptor transcription
bonds in collagen. factor that regulates drug metabolism and
Colibacillosis Infection with Escherichia coli. homoeostasis.
Colic A broad term which refers to episodes of Consumption Term used to describe wast-
uncontrollable, extended crying in a baby who ing of tissues including but not limited to
is otherwise healthy and well fed. tuberculosis.
Colitis Inflammatory bowel disease affecting Consumptive Afflicted with or associated with
the tissue that lines the gastrointestinal system. pulmonary tuberculosis.
Collyrium A lotion or liquid wash used as a Contraceptive An agent that reduces the likeli-
cleanser for the eyes, particularly in diseases hood of or prevents conception.
of the eye. Contraindication A condition which makes a
Colorectal Relating to the colon or rectum. particular treatment or procedure inadvisable.
296 Medical Glossary
Cryptococcal meningitis A fungal infection of category of signalling molecules that are used
the membranes covering the brain and spinal extensively in cellular communication.
cord (meninges). Cytopathic Any detectable, degenerative
Crytochidism (cryptochism) A developmen- changes in the host cell due to infection.
tal defect characterised by the failure of one Cytoprotective Protecting cells from noxious
or both testes to move into the scrotum as the chemicals or other stimuli.
male fetus develops. Cytosolic Relates to the fluid of the cytoplasm
Curettage Surgical procedure in which a body in cells.
cavity or tissue is scraped with a sharp instru- Cytostatic Preventing the growth and prolif-
ment or aspirated with a cannula. eration of cells.
Cutaneous Pertaining to the skin. Cytotoxic Of or relating to substances that are
CXC8 Also known as interleukin 8, IL-8. toxic to cells; cell killing.
Cyanogenesis Generation of cyanide. adj. d-Galactosamine An amino sugar with unique
cyanogenetic. hepatotoxic properties in animals.
Cyclooxygenase (COX) An enzyme that Dandruff Scurf, dead, scaly skin among the
is responsible for the formation of pros- hair.
tanoids—prostaglandins, prostacyclins and Dartre Condition of dry, scaly skin
thromboxanes that are each involved in the Debility Weakness, relaxation of muscular
inflammatory response. Two different COX fibre.
enzymes existed, now known as COX-1 and Debridement Is the process of removing non-
COX-2. living tissue from pressure ulcers, burns and
Cyclooxygenase-1 (COX-1) Is known to be other wounds.
present in most tissues. In the gastrointestinal Debriding agent Substance that cleans and
tract, COX-1 maintains the normal lining of treats certain types of wounds, burns and
the stomach. The enzyme is also involved in ulcers.
kidney and platelet function. Deciduogenic Relating to the uterus lining that
Cyclooxygenase-2 (COX-2) Is primarily pres- is shed off at childbirth.
ent at sites of inflammation. Deciduoma Decidual tissue induced in the
Cysteine proteases Are enzymes that degrade uterus (as by trauma) in the absence of
polypeptides possessing a common catalytic pregnancy.
mechanism that involves a nucleophilic cys- Deciduomata Plural of deciduoma.
teine thiol in a catalytic triad. They are found Decidual stromal cells Like endometrial
in fruits like papaya, pineapple and kiwifruit. glands and endothelium, express integrins that
Cystitis A common urinary tract infection that bind basement components.
occurs when bacteria travel up the urethra, Decoction A medical preparation made by
infect the urine and inflame the bladder lining. boiling the ingredients.
Cystorrhea Discharge of mucus from the Decongestant A substance that relieves or
bladder. reduces nasal or bronchial congestion.
Cytochrome bc-1 complex Ubihydroquinone: Deep venous thrombosis Is a blood clot that
Cytochrome c oxidoreductase. forms in a vein deep inside a part of the body.
Cytochrome P450 3A CYP3A A very large Defibrinated plasma Blood whose plasma
and diverse superfamily of haem-thiolate pro- component has had fibrinogen and fibrin
teins found in all domains of life. This group removed.
of enzymes catalyses many reactions involved Degranulation Cellular process that releases
in drug metabolism and synthesis of choles- antimicrobial cytotoxic molecules from secre-
terol, steroids and other lipids. tory vesicles called granules found inside
Cytokine Non-antibody proteins secreted by some cells.
certain cells of the immune system which Delayed after depolarisations
carry signals locally between cells. They are a (DADs) Abnormal depolarisation that
298 Medical Glossary
begins during phase 4—after depolarisation is tion of some mucocutaneous conditions such as
completed, but before another action potential lichen planus or the vesiculobullous disorders.
would normally occur. Detoxifier A substance that promotes the
Delirium Is common, sudden severe confusion removal of toxins from a system or organ.
and rapid changes in brain function that occur Diabetes A metabolic disorder associated with
with physical or mental illness; it is reversible inadequate secretion or utilisation of insulin
and temporary. and characterised by frequent urination and
Demulcent An agent that soothes internal persistent thirst. See Diabetes mellitus.
membranes. Also called emollient. Diabetes mellitus (DM) (Sometimes called
Dendritic cells Are immune cells and form ‘sugar diabetes’) is a set of chronic, meta-
part of the mammalian immune system, func- bolic disease conditions characterised by high
tioning as antigen-presenting cells. blood sugar (glucose) levels that result from
Dentition A term that describes all of the upper defects in insulin secretion, or action, or both.
and lower teeth collectively. Diabetes mellitus appears in two forms.
Deobstruent A medicine which removes Diabetes mellitus type I (Formerly known as
obstructions; also called an aperient. juvenile onset diabetes) caused by deficiency
Deoxypyridinoline (Dpd) A crosslink prod- of the pancreatic hormone insulin as a result
uct of collagen molecules found in bone and of destruction of insulin-producing beta cells
excreted in urine during bone degradation. of the pancreas. Lack of insulin causes an
Depilatory An agent for removing or destroy- increase of fasting blood glucose that begins to
ing hair. appear in the urine above the renal threshold.
Depressant A substance that diminishes func- Diabetes mellitus type II (Formerly called
tional activity, usually by depressing the ner- non-insulin-dependent diabetes mellitus or
vous system. adult-onset diabetes) the disorder is charac-
Depurative An agent used to cleanse or purify terised by high blood glucose in the context
the blood; it eliminates toxins and purifies the of insulin resistance and relative insulin defi-
system. ciency in which insulin is available but cannot
Dermatitis Inflammation of the skin causing be properly utilised.
discomfort such as eczema. Diabetic autonomic neuropathy (DAN) is a
Dermatitis herpetiformis An autoimmune serious and common complication of diabetes
chronic blistering skin disorder characterised involving damage of the autonomic nerves.
by blisters filled with a watery fluid. Major clinical manifestations of DAN include
Dermatophyte A fungus parasitic on the skin. resting tachycardia, exercise intolerance, ortho-
Dermatosis Is a broad term that refers to any static hypotension, constipation, gastroparesis,
disease of the skin, especially one that is not erectile dysfunction, sudomotor dysfunction,
accompanied by inflammation. impaired neurovascular function, ‘brittle dia-
Dermonecrotic Pertaining to or causing necro- betes’ and hypoglycaemic autonomic failure.
sis of the skin. Diabetic foot Any pathology that results
Dermopathy A skin disorder characterised by directly from diabetes mellitus or any long-
discoloured patches and small papules that term or chronic complication of diabetes
often become pigmented and ulcerated and mellitus.
result in scars, most commonly occurring on Diabetic neuropathy A neuropathic disor-
the shins of people with diabetes mellitus. der that is associated with diabetes mellitus.
Desmutagen Substances that inactivate muta- It affects all peripheral nerves including pain
gens (cancer-causing agents). fibres, motor neurons and the autonomic ner-
Desquamation The shedding of the outer lay- vous system.
ers of the skin. Diabetic retinopathy Damage to the retina
Desquamative ginginvitis Red, painful, glazed caused by complications of diabetes mellitus,
and friable gingivae which may be a manifesta- which can eventually lead to blindness.
Medical Glossary 299
Diads Two adjacent structural units in a poly- Distal sensory polyneuropathy (DSPN) Or
mer molecule. peripheral neuropathy, is the most common
Dialysis Is a method of removing toxic sub- neurological problem in HIV disease. DSPN
stances (impurities or wastes) from the blood also represents a complex symptom that
when the kidneys are unable to do so. occurs because of peripheral nerve damage
Diaphoresis Is profuse sweating commonly related to advanced HIV disease.
associated with shock and other medical Diuresis Increased urination.
emergency conditions. Diuretic A substance that increases urination
Diaphoretic A substance that induces perspira- (diuresis).
tion. Also called sudorific. Diverticular disease Is a condition affecting
Diaphyseal Pertaining to or affecting the shaft the large bowel or colon and is thought to be
of a long bone (diaphysis). caused by eating too little fibre.
Diaphysis The main or midsection (shaft) of a Diverticulitis Common, sometimes painful
long bone. digestive disease which involves the forma-
Diarrhoea A profuse, frequent and loose dis- tion of pouches (diverticula) within the bowel
charge from the bowels. wall.
Diastolic Referring to the time when the heart DMBA 7,12-Dimethylbenzanthracene. A poly-
is in a period of relaxation and dilatation cyclic aromatic hydrocarbon found in tobacco
(expansion). cf. systolic. smoke that is a potent carcinogen.
Dieresis Surgical separation of parts. DNA Deoxyribonucleic acid, a nucleic acid
Dietary fibre Is a term that refers to a group that contains the genetic instructions used in
of food components that pass through the the development and functioning of all known
stomach and small intestine undigested and living organisms.
reach the large intestine virtually unchanged. DOCA Desoxycorticosterone acetate—a ste-
Scientific evidence suggest that a diet high roid chemical used as replacement therapy in
in dietary fibre can be of value for treating Addison’s disease.
or preventing such disorders as constipation, Dopamine A catecholamine neurotransmit-
irritable bowel syndrome, diverticular disease, ter that occurs in a wide variety of animals,
hiatus hernia and haemorrhoids. Some com- including both vertebrates and invertebrates.
ponents of dietary fibre may also be of value Dopaminergic Relating to, or activated by the
in reducing the level of cholesterol in blood neurotransmitter, dopamine.
and thereby decreasing a risk factor for coro- Double blind Refers to a clinical trial or exper-
nary heart disease and the development of iment in which neither the subject nor the
gallstones. Dietary fibre is beneficial in the researcher knows which treatment any par-
treatment of some diabetics. ticular subject is receiving.
Digalactosyl diglycerides Are the major lipid Douche A localised spray of liquid directed
components of chloroplasts. into a body cavity or onto a part.
Diosgenin A steroid-like substance that is involved DPPH 2,2-Diphenyl-1-picryl-hydrazyl—a
in the production of the hormone progesterone, crystalline, stable free radical used as an
extracted from roots of Dioscorea yam. inhibitor of free radical reactions.
Dipsia Sensation of dryness in the mouth and Dracunculiasis Also called guinea worm dis-
throat related to a desire to drink. ease (GWD), is a parasitic infection caused by
Dipsomania Pathological use of alcohol. the nematode, Dracunculus medinensis.
Discutient An agent (as a medicinal applica- Dropsy An old term for the swelling of soft tis-
tion) which serves to disperse morbid matter. sues due to the accumulation of excess water.
Disinfectant An agent that prevents the spread adj. dropsical.
of infection, bacteria or communicable Drug metabolising enzymes Play central roles
disease. in the biotransformation, metabolism and/or
300 Medical Glossary
Eczematous rash Dry, scaly, itchy rash. Embryotoxic Term that describes any chemi-
ED 50 Is defined as the dose producing a cal which is harmful to an embryo.
response that is 50 % of the maximum Emesis Vomiting, throwing up.
obtainable. Emetic An agent that induces vomiting. cf.
Edema Formerly known as dropsy or hydropsy, antiemetic
is characterised as swelling caused by abnor- Emetocathartic Causing vomiting and
mal accumulation of fluid beneath the skin, or purging.
in one or more cavities of the body. It usually Emmenagogue A substance that stimulates,
occurs in the feet, ankles and legs, but it can initiates and/or promotes menstrual flow.
involve the entire body. Emmenagogues are used in herbal medicine
Edematogenic Producing or causing edema. to balance and restore the normal function of
EGFR proteins Epidermal growth factor the female reproductive system.
receptor (EGFR) proteins. Protein kinases are Emollient An agent that has a protective and
enzymes that transfer a phosphate group from soothing action on the surfaces of the skin and
a phosphate donor onto an acceptor amino membranes.
acid in a substrate protein. Emphysema A long-term, progressive disease
EGR-1 Early growth response 1, a human of the lungs that primarily causes shortness of
gene. breath.
Eicosanoids Are signalling molecules made Emulsion A preparation formed by the suspen-
by oxygenation of arachidonic acid, a twenty- sion of very finely divided oily or resinous liq-
carbon essential fatty acid, and include prosta- uid in another liquid.
glandins and related compounds. Encephalitis Inflammation of the brain caused
Elastase A serine protease that also hydrolyses by a virus.
amides and esters. Encephalocele A neural tube defect character-
Electrocardiography Or ECG, is a transtho- ised by saclike protrusions of the brain tissue
racic interpretation of the electrical activity through a congenital fissure in the skull.
of the heart over time captured and externally Encephalomalacia Cerebral softening, a
recorded by skin electrodes. localised softening of the brain substance, due
Electromyogram (EMG) A test used to to haemorrhage or inflammation.
record the electrical activity of muscles. Encephalopathy A disorder or disease of the
An electromyogram (EMG) is also called a brain.
myogram. Endocrine adj. of or relating to endocrine
Electuary A medicinal paste composed of glands or the hormones secreted by them.
powders, or other medical ingredients, incor- Endocytosis Is the process by which cells
porated with sweeteners to hide the taste, suit- absorb material (molecules such as proteins)
able for oral administration. from outside the cell by engulfing it with their
Elephantiasis A disorder characterised by cell membrane.
chronic thickened and oedematous tissue on Endometrial cancer Cancer that arises in the
the genitals and legs due to various causes. endometrium, the lining of the uterus (womb).
11β-Hydroxysteroid dehydrogenase (HSD- Endometriosis Is a common and often painful
11β or 11β-HSD) is the name of a family of disorder of the female reproductive system in
enzymes that catalyse the conversion of inert which the endometrium, the tissue that nor-
11 keto-products (cortisone) to active cortisol, mally lines the womb (uterus), grows outside
or vice versa. the uterus. The two most common symptoms
Embrocation Lotion or liniment that relieves of endometriosis are pain and infertility.
muscle or joint pains. Endometritis Refers to inflammation of the
Embryonation Formation of embryo in the endometrium, the inner lining of the uterus.
egg. Endometrium The inner lining of the uterus.
302 Medical Glossary
Epileptiform Resembling epilepsy or its mani- tidylserine exposure at the outer membrane
festations. adj. epileptiformic. leaflet.
Epileptogenesis A process by which a normal Erysipelas Is an intensely red Streptococcus
brain develops epilepsy, a chronic condition in bacterial infection that occurs on the face and
which seizures occur. adj. epileptogenic. lower extremities.
Episiotomy A surgical incision through the Erythema Abnormal redness and inflamma-
perineum made to enlarge the vagina and tion of the skin, due to vasodilation.
assist childbirth. Erythema multiforme Is a skin disorder
Epithelial–mesenchymal transition or trans- due to an allergic reaction or infection;
formation (EMT) A process by which characterised by fever, general ill feeling,
epithelial cells lose their cell polarity and cell– skin itching, joint aches and multiple skin
cell adhesion and gain migratory and invasive lesions.
properties to become mesenchymal cells. Erythematous Characterised by erythema.
Epithelioma A usually benign skin disease Erythroderma Exfoliative dermatitis.
most commonly occurring on the face, around Erythroleukoplakia An abnormal patch of
the eyelids and on the scalp. red and white tissue that forms on mucous
Epitope A single antigenic site on a protein membranes in the mouth and may become
against which an antibody reacts. cancer. Tobacco (smoking and chew-
Epitrochlearis The superficial-most muscle of ing) and alcohol may increase the risk of
the arm anterior surface. erythroleukoplakia.
Epistaxis Acute haemorrhage from the nostril, Erythropoiesis Is the process whereby ery-
nasal cavity or nasopharynx (nosebleed). throid precursor cells proliferate and differen-
Epstein–Barr virus Herpesvirus that is the tiate into red blood cells.
causative agent of infectious mononucleo- Erythropoietin (EPO) A hormone produced
sis. It is also associated with various types of by the kidney that promotes the formation
human cancers. of red blood cells (erythrocytes) in the bone
ERbeta Oestrogen receptor beta, a nuclear marrow.
receptor which is activated by the sex hor- Eschar A slough or piece of dead tissue that is
mone, oestrogen. cast off from the surface of the skin.
Ergocalciferol A form of vitamin D, also Escharotic Capable of producing an eschar; a
called vitamin D2. See Vitamin D. caustic or corrosive agent.
Ergogenic Increasing capacity for bodily Estradiol Is the predominant sex hormone
or mental labour, especially by eliminating present in females, also called oestradiol.
fatigue symptoms. Estrogenic Relating to oestrogen or producing
Ergonic Increasing capacity for bodily or men- oestrus.
tal labour, especially by eliminating fatigue Euglycaemia Normal blood glucose
symptoms. concentration.
ERK (extracellular signal regulated Eupeptic Conducive to digestion.
kinases) Widely expressed protein kinase Exanthema Sudden widespread rash.
intracellular signalling molecules which are Exanthematous Characterised by or of the
involved in functions including the regulation nature of an eruption or rash.
of meiosis, mitosis and postmitotic functions Excitotoxicity Is the pathological process by
in differentiated cells. which neurons are damaged and killed by glu-
Eructation The act of belching or of casting up tamate and similar substances.
wind from the stomach through the mouth. Excipient A pharmacologically inert substance
Eruption A visible rash or cutaneous used as a diluent or vehicle for the active
disruption. ingredients of a medication.
Eryptosis Suicidal death of erythrocytes, Exencephaly A type of cephalic disorder
characterised by cell shrinkage, membrane wherein the brain is located outside of the
blebbing, activation of proteases and phospha- skull.
304 Medical Glossary
Fibroblast Type of cell that synthesises the intolerance, certain foods or a sudden switch
extracellular matrix and collagen, the struc- to a high fibre.
tural framework (stroma) for animal tissues, Flavans A subgroup of flavonoids. See
and plays a critical role in wound healing. Flavonoids.
Fibrogenic Promoting the development of Flavanols A subgroup of flavonoids, are a
fibres. class of flavonoids that use the 2-phenyl-3,4-
Fibromyalgia A common and complex dihydro-2H-chromen-3-ol skeleton. These
chronic, body-wide pain disorder that affects compounds include the catechins and the cat-
people physically, mentally and socially. echin gallates. They are found in chocolate,
Symptoms include debilitating fatigue, sleep fruits and vegetables. See Flavonoids.
disturbance and joint stiffness. Also referred Flavanones A subgroup of flavonoids, consti-
to as FM or FMS. tute >90% of total flavonoids in citrus. The
Fibronectin A high molecular weight major dietary flavanones are hesperetin, nar-
(~440kDa) glycoprotein of the extracellular ingenin and eriodictyol.
matrix (ECM) that adheres to membrane- Flavivirus A family of viruses transmitted by
spanning receptor proteins called integrins. mosquitoes and ticks that cause some impor-
Fibrosarcoma A malignant tumour derived tant diseases, including dengue, yellow fever,
from fibrous connective tissue and character- tick-borne encephalitis and West Nile fever.
ised by immature proliferating fibroblasts or Flavones A subgroup of flavonoids based on
undifferentiated anaplastic spindle cells. the backbone of 2-phenylchromen-4-one
Fibrosis The formation of fibrous tissue as a (2-phenyl-1-benzopyran-4-one). Flavones are
reparative or reactive process. mainly found in cereals and herbs.
Filarial Pertaining to a thread-like nematode Flavonoids (Or bioflavonoids) are a group of
worm. polyphenolic antioxidant compounds that
Filariasis A parasitic and infectious tropical dis- occur in plant as secondary metabolites. They
ease that is caused by thread-like filarial nema- are responsible for the colour of fruit and veg-
tode worms in the superfamily Filarioidea. etables. Twelve basic classes (chemical types)
Fistula An abnormal connection between two of flavonoids have been recognised: flavones,
organs inside of the body. isoflavones, flavans, flavanones, flavanols, fla-
Fistula-in-ano A track connecting the internal vanolols, anthocyanidins, catechins (includ-
anal canal to the skin surrounding the anal ing proanthocyanidins), leucoanthocyanidins,
orifice. chalcones, dihydrochalcones and aurones.
5′-Nucleotidase (5′-Ribonucleotide phospho- Apart from their antioxidant activity, flavo-
hydrolase) an intrinsic membrane glycoprotein noids are known for their ability to strengthen
present as an ectoenzyme in a wide variety of capillary walls, thus assisting circulation and
mammalian cells, hydrolyses 5′-nucleotides helping to prevent and treat bruising, vari-
to their corresponding nucleosides. cose veins, bleeding gums and nosebleeds
5-HT1A receptor A serotonin protein that and heavy menstrual bleeding, and are also
binds to 5-5-hydroxytryptamine (5-HT) with anti-inflammatory.
high affinity to exert subtle control over emo- Flourine F is an essential chemical element
tion and behaviour. that is required for the maintenance of healthy
Flash electroretinogram or flash ERG (fERG bones and teeth and to reduce tooth decay. It
) Is a test which measures the electrical is found in seaweeds, tea, water, seafood and
response of the eye’s light-sensitive cells (rods dairy products.
and cones). It also checks other cell layers in Fluorosis A dental health condition caused by
the retina. a child receiving too much fluoride during
Flatulence Is the presence of a mixture of tooth development.
gases known as flatus in the digestive tract Flux An excessive discharge of fluid.
of mammals expelled from the rectum. FMD (flow-mediated dilation) A measure of
Excessive flatulence can be caused by lactose endothelial dysfunction which is used to eval-
306 Medical Glossary
uate cardiovascular risk. Also called FMVD ( Staphylococcus aureus, resulting in the local-
flow-mediated vasodilation). ised accumulation of pus and dead tissue.
Focal adhesion kinase (FAK) Is a protein Furunculosis Skin condition characterised by
tyrosine kinase which is recruited at an early persistent, recurring boils.
stage to focal adhesions and which mediates G protein-coupled receptor kinases (GRKs,
many of the downstream regulatory responses. GPCRKs) A family of protein kinases which
Follicle-stimulating hormone (FSH) A hor- regulate the activity of G protein-coupled
mone produced by the pituitary gland. In receptors (GPCRs) by phosphorylating their
women, it helps control the menstrual cycle intracellular domains after their associated G
and the production of eggs by the ovaries. proteins have been released and activated.
Follicular atresia The breakdown of the ovar- GABA Gamma aminobutyric acid, required
ian follicles. as an inhibitory neurotransmitter to block the
Fomentation Treatment by the application of transmission of an impulse from one cell to
war, moist substance. another in the central nervous system, which
Fontanelle Soft spot on an infant’s skull. prevents over-firing of the nerve cells. It is
Forkhead box-O transcription factors used to treat both epilepsy and hypertension.
(FOXOs) Are a family of transcription fac- GADD 152 A proapoptotic gene.
tors that play important roles in regulating the Galctifuge Or lactifuge, causing the arrest of
expression of genes involved in cell growth, milk secretion.
proliferation, differentiation and longevity. It Galactogogue A substance that promotes the
also plays an important role in tumour sup- flow of milk.
pression by regulating the expression of genes Galactophoritis Inflammation of the milk
involved in stress resistance, DNA damage ducts.
repair, cell cycle arrest and apoptosis. Galactopoietic Increasing the flow of milk;
Framboesia See Yaws. milk producing.
FRAP Ferric reducing ability of plasma, an Gallbladder A small, pear-shaped muscular
assay used to assess antioxidant property. sac, located under the right lobe of the liver, in
Fibrillation Is the rapid, irregular and unsyn- which bile secreted by the liver is stored until
chronised contraction of muscle fibres, espe- needed by the body for digestion. Also called
cially with regard to the heart. cholecyst, cholecystitis.
5-Dihydroaldosterone A hormone secreted Gallic acid equivalent (GAE) Measures the
by the adrenal cortex that regulates electro- total phenol content in terms of the standard
lyte and water balance by increasing the renal gallic acid by the Folin–Ciocalteu assay.
retention. Galphai proteins or G alpha I proteins Are
Friedreich’s ataxia Is a genetic inherited dis- heterotrimeric guanine nucleotide-regulatory
order that causes progressive damage to the (G) proteins associated with a variety of intra-
nervous system resulting in symptoms rang- cellular membranes and specific plasma mem-
ing from muscle weakness and speech prob- brane domains.
lems to heart disease. cf. ataxia. Gamma GT (GGT) Gamma-glutamyl trans-
Fulminant hepatitis Acute liver failure. peptidase, a liver enzyme.
Functional dyspepsia A non-ulcer condition Gap junction intercellular communication Is
that causes an upset stomach or pain or dis- considered to be the sole means by which low
comfort in the upper belly, near the ribs. molecular weight factors inside a cell can pass
Functional food Is any fresh or processed directly into the interior of neighbouring cells.
food claimed to have a health-promoting or Gap junctions are considered to play an essen-
disease-preventing property beyond the basic tial role in the maintenance of homeostasis.
function of supplying nutrients. Also called Gastralgia (Heartburn) pain in the stomach
medicinal food. or abdominal region. It is caused by excess
Furuncle Is a skin disease caused by the of acid, or an accumulation of gas, in the
infection of hair follicles usually caused by stomach.
Medical Glossary 307
Gastric Pertaining to or affecting the stomach. Glaucoma A group of eye diseases in which
Gastric emptying Refers to the speed at which the optic nerve at the back of the eye is slowly
food and drink leave the stomach. destroyed, leading to impaired vision and
Gastritis Inflammation of the stomach. blindness.
Gastrocnemius muscle The big calf muscle at Gleet A chronic inflammation (as gonorrhoea)
the rear of the lower leg. of a bodily orifice usually accompanied by an
Gastrodynia Pain in the stomach. abnormal discharge.
Gastroparesis Also called delayed gastric Glial cells Support non-neuronal cells in the
emptying, a medical condition consisting of central nervous system that maintain homeo-
a paresis (partial paralysis) of the stomach, stasis, form myelin and provide protection for
resulting in food remaining in the stomach for the brain’s neurons.
an abnormally long time. Glioma Is a type of tumour that starts in the
Gastroprokinetic See Prokinetic. brain or spine. It is called a glioma because it
Gastrotonic (gastroprotective) Substance that arises from glial cells.
strengthens, tones or regulates gastric func- Glioblastoma Common and most lethal form
tions (or protects from injury) without overt of brain tumour.
stimulation or depression. Glioblastoma multiforme Most common and
Gavage Forced feeding. most aggressive type of primary brain tumour
Gene silencing Suppression of the expression in humans, involving glial cells.
of a gene. Glomerulonephritis (GN) A renal disease
Genotoxic Describes a poisonous substance characterised by inflammation of the glom-
which harms an organism by damaging its eruli, or small blood vessels in the kidneys.
DNA, thereby capable of causing mutations Also known as glomerular nephritis. adj.
or cancer. glomerulonephritic.
Genotoxin A chemical or another agent that Glomerulopathy Any disease of the renal
damages cellular DNA, resulting in mutations glomeruli.
or cancer. Glomerulosclerosis A hardening (fibrosis) of
Geriatrics Is a subspecialty of internal medicine the glomerulus in the kidney.
that focuses on healthcare of elderly people. Glossal Pertaining to the tongue.
Gestational hypertension Development of GLP-1 Glucagon-like peptide-1.
arterial hypertension in a pregnant woman Glucagon-like peptide-1 (GLP-1) is derived
after 20 weeks of gestation. from the transcription product of the proglu-
Ghrelin A gastrointestinal peptide hormone cagon gene, reduces insulin requirement in
secreted by epithelial cells in the stomach lin- diabetes mellitus and promotes satiety.
ing; it stimulates appetite and gastric empty- Gluconeogenesis A metabolic pathway that
ing and increases cardiac output. results in the generation of glucose from non-
Gingival index An index describing the clini- carbohydrate carbon substrates such as lac-
cal severity of gingival inflammation as well tate. adj. gluconeogenic.
as its location. Glucose-6-phosphate dehydrogenase (G6PD
Gingivitis Refers to gingival inflammation or G6PDH) Is a cytosolic enzyme in the
induced by bacterial biofilms (also called pentose phosphate metabolic pathway.
plaque) adherent to tooth surfaces. Glucose transporter type 4 (GLUT 4) Insulin-
Gin-nan sitotoxism Toxicity caused by inges- regulated glucose transporter found in adipose
tion of ginkgotoxin and characterised mainly tissues and striated muscles that modulates
by epileptic convulsions, paralysis of the legs insulin-related translocation into the cell.
and loss of consciousness. Glucose transporters (GLUT or SLC2A fam-
GIP Gastric inhibitory polypeptide also known ily) are a family of membrane proteins found
as the glucose-dependent insulinotropic in most mammalian cells.
peptide, a member of the secretin family of Glucosuria or glycosuria Is the excretion of
hormones. glucose into the urine.
308 Medical Glossary
in the GSH/GSSG ratio are often used to inflammatory conditions by oxidant stress, an
assess exposure of cells to oxidative stress. enzyme that catalyses degradation of haem.
GSTM Glutathione S transferase M1, a major Haemochromatosis iron overload in the body
group of detoxification enzymes. with a hereditary or primary cause.
GSTM 2 Glutathione S transferase M2, a Haemodialysis, hemodialysis A method for
major group of detoxification enzymes. removing waste products such as potassium
G2-M cell cycle The phase where the cell pre- and urea, as well as free water from the blood
pares for mitosis and where chromatids and when the kidneys are in renal failure.
daughter cells separate. Haemolyis Lysis of red blood cells and the
Guillain–Barre syndrome Is a serious dis- release of haemoglobin into the surrounding
order that occurs when the body’s defence fluid (plasma). adj. haemolytic.
(immune) system mistakenly attacks part of Haemoptysis, hemoptysis Is the coughing up
the nervous system, leading to nerve inflam- of blood from the respiratory tract. The blood
mation, muscle weakness and other symptoms. can come from the nose, mouth, throat and the
Gynecomastia Enlargement of the gland tissue airway passages leading to the lungs.
of the male breast, resulting from an imbal- Haemorrhage, hemaorrhage bleeding, dis-
ance of hormones. charge of blood from blood vessels.
Gynecopathy Any or various diseases specific Haemorrhoids, hemorrhoids A painful con-
to women. dition in which the veins around the anus
Gynoid adiposity Fat distribution mainly to or lower rectum are enlarged, swollen and
the hips and thighs, pear shaped. inflamed. Also called piles.
Haemagogic Promoting a flow of blood. Haemostasis, hemostasis A complex process
Haematemesis, hematemesis Is the vomiting which causes the bleeding process to stop.
of blood. Haemostatic, hemostatic Something that
Haematinic Improving the quality of the stops bleeding.
blood, its haemoglobin level and the number Halitosis (Bad breath) a common condition
of erythrocytes. caused by sulphur-producing bacteria that live
Haematochezia Passage of stools containing within the surface of the tongue and in the
blood. throat.
Haematochyluria, hematochyluria the dis- Hallucinogen Drug that produces hallucinogen.
charge of blood and chyle (emulsified fat) in Hallucinogenic Inducing hallucinations.
the urine; see also Chyluria. Hallux abducto valgus Commonly called
Haematoma, hematoma A localised accumu- bunion is an abnormal bending of the big toe
lation of blood in a tissue or space composed towards the other toes of the foot.
of clotted blood. Haplotype A set of alleles of closely linked
Haematometra, hematometra A medical loci on a chromosome that tend to be inherited
condition involving bleeding of or near the together.
uterus. Hapten A small molecule that can elicit an
Haematopoiesis, hematopoiesis Formation of immune response only when attached to a
blood cellular components from the haemato- large carrier such as a protein.
poietic stem cells. HATs Histone acetyl transferases, enzymes
Haematopoietic adj. relating to the formation that regulate the acetylation of histones and
and development of blood cells. transcription factors, playing a major role in
Haematuria, Hematuria Is the presence of the growth and differentiation of cells.
blood in the urine. Haematuria is a sign that HbA1c Glycosylated haemoglobin.
something is causing abnormal bleeding in a HBeAg Hepatitis B e antigen.
person’s genitourinary tract. HBsAg Hepatitis B s antigen.
Haeme oxygenase (HO-1, encoded by Hmox1) HBD-2 (human β-defensin 2) A member of
is an inducible protein activated in systemic the defensin family of antimicrobial peptides
310 Medical Glossary
that plays important roles in the innate and Hemorheology Study of blood flow and
adaptive immune system of both vertebrates its elements in the circulatory system. adj.
and invertebrates. hemorheological.
Heartburn Burning sensation in the stomach Haemorrhagic colitis An acute gastroenteritis
and oesophagus caused by excessive acidity characterised by overtly bloody diarrhoea that
of the stomach fluids. is caused by Escherichia coli infection.
Heat rash Any condition aggravated by heat or Haemolysin Certain proteins and lipids that
hot weather such as intertrigo. cause lysis of red blood cells by damaging
Heat shock chaperones (HSC) Ubiquitous their cell membranes.
molecules involved in the modulation of pro- Haemolytic uremic syndrome Is a disease
tein conformational and complexation states, characterised by haemolytic anaemia, acute
associated with heat stress or other cellular renal failure (uraemia) and a low platelet
stress responses. count.
Heat shock proteins ( HSP) A group of func- Hepa-1c1c7 A type of hepatoma cells.
tionally related proteins, the expression of Hepatalgia Pain or discomfort in the liver area.
which is increased when the cells are exposed Hepatectomy The surgical removal of part or
to elevated temperatures or other cellular all of the liver.
stresses. Hepatic Relating to the liver.
Helminthiasis A disease in which a part of the Hepatic cirrhosis Affecting the liver, charac-
body is infested with worms such as pinworm, terised by hepatic fibrosis and regenerative
roundworm or tapeworm. nodules.
Haemagglutination A specific form of agglu- Hepatic encephalopathy Is the loss of brain
tination that involves red blood cells. function that occurs when the liver is unable
Haemagglutination–inhibition test Measures to remove toxins from the blood.
the ability of soluble antigen to inhibit the Hepatic fibrosis Is overly profuse wound
agglutination of antigen-coated red blood healing in which excessive connective tissue
cells by antibodies. builds up in the liver.
Haemagglutinin Refers to a substance that Hepatitis Inflammation of the liver.
causes red blood cells to agglutinate. Hepatitis A (Formerly known as infectious
Haemangioma Blood vessel. hepatitis) is an acute infectious disease of the
Haematocrit Is a blood test that measures the liver caused by the hepatovirus hepatitis A
percentage of the volume of the whole blood virus.
that is made up of red blood cells. Hepatocarcinogenesis Represents a linear and
Haematopoietic Pertaining to the formation of progressive cancerous process in the liver in
blood or blood cells. which successively more aberrant monoclonal
Haematopoietic stem cell Is a cell isolated populations of hepatocytes evolve.
from the blood or bone marrow that can renew Hepatocellular carcinoma (HCC) Also called
itself and can differentiate to a variety of spe- malignant hepatoma, is a primary malignancy
cialised cells. (cancer) of the liver.
Haem oxygenase-1 (HO-1) An enzyme that Hepatocytolysis Cytotoxicity (dissolution) of
catalyses the degradation of haem; an induc- liver cells.
ible stress protein, confers cytoprotection Hepatoma Cancer of the liver.
against oxidative stress in-vitro and in-vivo. Hepatomegaly Condition of enlarged liver.
Hemiplegia Paralysis of the arm, leg and trunk Hepatopathy A disease or disorder of the liver.
on the same side of the body. Hepatoprotective (Liver protector) a sub-
Haemoglobinopathies Genetic defects that stance that helps protect the liver from dam-
produce abnormal haemoglobins and anaemia. age by toxins, chemicals or other disease
Haemolytic anaemia Anaemia due to haemol- processes.
ysis, the breakdown of red blood cells in the Hepatoregenerative A compound that pro-
blood vessels or elsewhere in the body. motes hepatocellular regeneration, repairs
Medical Glossary 311
and restores liver function to optimum High-density lipoprotein (HDL) Is one of the
performance. five major groups of lipoproteins which enable
Hepatotonic (Liver tonic ) a substance that is cholesterol and triglycerides to be transported
tonic to the liver—usually employed to nor- within the water-based blood stream. HDL
malise liver enzymes and function. can remove cholesterol from atheroma within
Hernia Occurs when part of an internal organ arteries and transport it back to the liver for
bulges through a weak area of muscle. excretion or reutilisation—which is the main
HER-2 Human epidermal growth factor recep- reason why HDL-bound cholesterol is some-
tor 2, a protein giving higher aggressiveness in times called ‘good cholesterol’, or HDL-C. A
breast cancer, also known as ErbB-2, ERBB2. high level of HDL-C seems to protect against
Herpes A chronic inflammation of the skin or cardiovascular diseases. cf. LDL.
mucous membrane characterised by the devel- HGPRT, HPRT (hypoxanthine-guanine phos-
opment of vesicles on an inflammatory base. phoribosyl transferase) An enzyme that
Herpes circinatus Dermatitis herpetiformis catalyses the conversion of 5-phosphoribosyl
(resembling herpes). -1-pyrophosphate and hypoxanthine, guanine
Herpes simplex virus 1 and 2 (HSV-1 and HSV- or 6-mercaptopurine to the corresponding
2) Are two species of the herpesvirus family 5′-mononucleotides and pyrophosphate. The
which cause a variety of illnesses/infections enzyme is important in purine biosynthesis as
in humans such as cold sores, chickenpox or well as central nervous system functions.
varicella, shingles or herpes zoster (VZV), Hippocampus A ridge in the floor of each lat-
cytomegalovirus (CMV) and various cancers eral ventricle of the brain that consists mainly
and can cause brain inflammation (encephali- of grey matter.
tis). HSV-1 is commonly associated with her- Hippocampal Pertaining to the hippocampus.
pes outbreaks of the face known as cold sores Hirsutism A condition where women have
or fever blisters, whereas HSV-2 is more often excess facial and body hair that is dark and
associated with genital herpes. They are also coarse.
called human herpesvirus 1 and 2 (HHV-1 and Histaminergic Liberated or activated by his-
HHV-2) and are neurotropic and neuroinva- tamine, relating to the effects of histamine at
sive viruses; they enter and hide in the human histamine receptors of target tissues.
nervous system, accounting for their durabil- Histaminergic receptors Are types of G
ity in the human body. protein-coupled receptors with histamine as
Herpes zoster Or simply zoster, commonly their endogenous ligand.
known as shingles and also known as zona, is Histone acetyltransferases (HAT) Are
a viral disease characterised by a painful skin enzymes that acetylate conserved lysine
rash with blisters. amino acids on histone proteins by transfer-
Herpes zoster ophthalmicus (HZO) Is a viral ring an acetyl group from acetyl CoA to form
ocular disease characterised by a painful skin e-N-acetyl lysine. HATs act as transcriptional
rash in one or more dermatome distributions coactivators.
of the fifth cranial nerve, shared by the eye and Histone lysine demethylases
orbit. (KDMs) Enzymes that play a key role in the
Heterophobia Term used to describe irrational amplification of hypoxia-inducible-factor sig-
fear of, aversion to, or discrimination against nalling and expression of proangiogenic genes
heterosexuals. in cancer and neurological disorders.
HDL-C (HDL cholesterol) High-density lipo- HIV See Human immunodeficiency virus.
protein cholesterol, also called ‘good choles- Hives (Urticaria) is a skin rash characterised by
terol’. See also High-density lipoprotein. circular wheals of reddened and itching skin.
Hiatus hernia Occurs when the upper part of HLA Human leucocyte antigen system, name
the stomach pushes its way through a tear in of the major histocompatibility complex
the diaphragm. (MHC) in humans.
312 Medical Glossary
HLA-DQB1 Human leucocyte antigen beta HSP70 Heat shock protein chaperone that con-
chain. fers protection against heat-induced apoptosis.
HLA-DR A major histocompatibility complex HSP90 A 90kDa heat shock protein chaperone
(MHC) class II cell surface receptor encoded that has the ability to regulate a specific subset
by the human leucocyte antigen complex on of cellular signalling proteins that have been
chromosome 6 region 6p21.31. implicated in disease processes.
HMG-CoAr 3-Hydroxy-3-methyl-glutaryl-- HSPD 1 Heat shock 60kDa protein 1
CoA reductase or HMGCR is the rate- hTERT (TERT) Telomerase reverse transcrip-
controlling enzyme (EC 1.1.1.88) of the tase is a catalytic subunit of the enzyme telom-
mevalonate pathway. erase in humans. It exerts a novel protective
HMG-CoA 3-Hydroxy-3-methylglutaryl- function by binding to mitochondrial DNA,
coenzyme A, an intermediate in the mevalon- increasing respiratory chain activity and
ate pathway. protecting against oxidative stress-induced
Hodgkin’s disease Disease characterised by damage.
enlargement of the lymph glands, spleen and HT29 cells Are human intestinal epithelial
anaemia. cells which produce the secretory component
Homeodomain transcription factor A protein of immunoglobulin A (IgA) and carcinoem-
domain encoded by a homeobox. Homeobox bryonic antigen (CEA).
genes encode transcription factors which typi- Human cytomegalovirus (HCMV) A DNA
cally switch on cascades of other genes. herpesvirus which is the leading cause
Homeostasis The maintenance of a constant of congenital viral infection and mental
internal environment of a cell or an organism, retardation.
despite fluctuations in the external. Human factor X A coagulation factor also
Homeotherapy Treatment or prevention of known by the eponym Stuart–Prower fac-
disease with a substance similar but not iden- tor or as thrombokinase, is an enzyme
tical to the causative agent of the disease. involved in blood coagulation. It synthe-
Homocysteine An amino acid in the blood. sised in the liver and requires vitamin K for
Homograft See Allograft. its synthesis.
Hormesis a Term used by toxicologists to refer Human immunodeficiency virus (HIV) A
to a biphasic dose response to an environmen- retrovirus that can lead to acquired immuno-
tal agent characterised by a low-dose stimu- deficiency syndrome (AIDS), a condition in
lation or beneficial effect and a high-dose humans in which the immune system begins
inhibitory or toxic effect. to fail, leading to life-threatening opportunis-
Hormonal (female) Substance that has a hor- tic infections.
mone-like effect similar to that of oestrogen Humoral immune response (HIR) Is the
and/or a substance used to normalise female aspect of immunity that is mediated by
hormone levels. secreted antibodies (as opposed to cell-
Hormonal (male) Substance that has a mediated immunity, which involves T lym-
hormone-like effect similar to that of testos- phocytes) produced in the cells of the B
terone and/or a substance used to normalise lymphocyte lineage (B cell).
male hormone levels. HUVEC Human umbilical vein endothelial
HRT Hormone replacement therapy, the cells.
administration of the female hormones, oes- Hyaluronidase Enzymes that catalyse the
trogen and progesterone and sometimes hydrolysis of certain complex carbohydrates
testosterone. like hyaluronic acid and chondroitin sulphates.
HSF-1 factor Major regulator of heat shock Hydatidiform A rare mass or growth that
protein transcription in eukaryotes. forms inside the uterus at the beginning of a
HSP27 Is an ATP-independent, 27kDa heat pregnancy.
shock protein chaperone that confers protec- Hydrocele Abnormal accumulation of fluid
tion against apoptosis. inside the scrotum.
Medical Glossary 313
Hypertonia Abnormal increase in muscle ten- low levels of parathyroid hormone (PTH).
sion and a reduced ability of the muscle to PTH plays a key role in modulating the bal-
stretch. ance of calcium and phosphorus levels in the
Hypertriglyceridaemia or hypertriglycae- body.
mia A disorder that causes high triglycer- Hypoperfusion Decreased blood flow through
ides in the blood. an organ, characterised by an imbalance
Hypertrophy Enlargement or overgrowth of of oxygen demand and oxygen delivery to
an organ. tissues.
Hyperuricaemia Is a condition characterised Hypophagic Undereating.
by abnormally high level of uric acid in the Hypophysectomy The surgical removal of the
blood. hypophysis (pituitary gland).
Hypoadiponectinemia The state of having too Hypospadias An abnormal birth defect in
low level of adiponectin, a major metabolic males in which the urethra opens on the under
endocrine, responsible for regulating things surface of the penis.
like glucose uptake and lipolysis (the break- Hypotensive Characterised by or causing
down of fat deposits); low adiponectin is a risk diminished tension or pressure, as abnormally
factor for both type II diabetes and metabolic low blood pressure.
syndrome. Hypothermia A condition in which an organ-
Hypoalbuminemia A medical condition ism’s temperature drops below that required
where levels of albumin in blood serum are for normal metabolism and body functions.
abnormally low. Hypothermic Relating to hypothermia, with
Hypocalcaemic tetany A disease caused by an subnormal body temperature.
abnormally low level of calcium in the blood Hypoxaemia Is the reduction of oxygen spe-
and characterised by hyperexcitability of the cifically in the blood.
neuromuscular system and results in carpo- Hypoxia A shortage of oxygen in the body.
pedal spasms. adj. hypoxic.
Hypochlorhydria Refer to states where the Hypoxia-inducible factors
production of gastric acid in the stomach is (HIFs) Transcription factors that respond
absent or low. to changes in available oxygen in the cellu-
Hypocholesterolemic (Cholesterol reducer) a lar environment, specifically, to deficiency in
substance that lowers blood cholesterol levels. oxygen.
Hypocitraturia Low amount of citrate in the ICAM-1 (inter-cellular adhesion molecule
urine, an important risk factor for kidney stone 1) Also known as CD54 (cluster of differ-
formation. entiation 54), is a protein that in humans is
Hypocorticism See Addison’s disease. encoded by the ICAM1 gene.
Hypocortisolism See Addison’s disease. IC50 The median maximal inhibitory concen-
Hypoesthesia (Or hypesthesia) refers to a tration; a measure of the effectiveness of a
reduced sense of touch or sensation, or a par- compound in inhibiting biological or bio-
tial loss of sensitivity to sensory stimuli. chemical function.
Hypoglycaemic An agent that lowers the con- I.C.V. (Intra-cerebroventricular) injection of
centration of glucose (sugar) in the blood. chemical into the right lateral ventricle of the
Hypogonadism syndrome Characterised by brain.
defects of the gonads, a diminished functional Iceterus Jaundice, yellowish pigmentation of
activity of the gonads—the testes and ovaries the skin.
in males and females, respectively. Ichthyosis Dry, rectangular, fishlike scales on
Hypokalemia Medical condition in which the the skin.
concentration of potassium (K+) in the blood Ichthyotoxic A substance which is poisonous
is low. to fish.
Hypoparathyroidism An uncommon condi- Icteric hepatitis An infectious syndrome of
tion in which your body secretes abnormally hepatitis characterised by jaundice, nausea,
Medical Glossary 315
Intermittent claudication An aching, crampy, also essential for the regulation of cell growth
tired and sometimes burning pain in the legs and differentiation. A deficiency of iron limits
that comes and goes, caused by peripheral oxygen delivery to cells, resulting in fatigue,
vascular disease. It usually occurs when walk- poor work performance and decreased immu-
ing and disappears after rest. nity. Conversely, excess amounts of iron can
Interoceptive Relating to stimuli arising from result in toxicity and even death. Dietary
within the body. sources include certain cereals, dark green
Interstitium The space between cells in a leafy vegetables, dried fruit, legumes, sea-
tissue. food, poultry and meat.
Interstitial Pertaining to the interstitium. Ischemia An insufficient supply of blood to an
Intertrigo An inflammation (rash) caused by organ, usually due to a blocked artery.
microbial infection in skin folds. Ischuria Retention or suppression of urine.
Intima Innermost layer of an artery or vein. Isoflavones A subgroup of flavonoids in which
Intimal hyperplasia The thickening of the the basic structure is a 3-phenyl chromane
tunica intima of a blood vessel as a complica- skeleton. They act as phytoestrogens in mam-
tion of a reconstruction procedure. mals. See Flavonoids.
Intoxicant Substance that produces drunken- Isomers Substances that are composed of the
ness or intoxication. same elements in the same proportions and
Intracavernosal Within the copus cavern- hence have the same molecular formula but
soum, columns of erectile tissues forming the differ in properties because of differences in
body of the penis. the arrangement of atoms.
Intraperitoneal (i.p.) The term used when Isoprostanes Unique prostaglandin-like com-
a chemical is contained within or adminis- pounds generated in-vivo from the free radi-
tered through the peritoneum (the thin, trans- cal-catalysed peroxidation of essential fatty
parent membrane that lines the walls of the acids.
abdomen). Jamu Traditional Indonesian herbal medicine.
Intrathecal (i.t.) Through the theca of the spi- Janus kinase (JAK)-signal transducer and
nal cord into the subarachnoid space. activator of transcription (STAT) signal-
Intromission The act of putting one thing into ling Are essential molecules in cytokine sig-
another. nal transduction pathways involved in cancer
Intubation Refers to the placement of a tube development and progression.
into an external or internal orifice of the body. Jaundice Refers to the yellow colour of the
Iodine (I) Is an essential chemical element skin and whites of the eyes caused by excess
that is important for hormone development bilirubin in the blood.
in the human body. Lack of iodine can lead JNK (Jun N-terminal kinase), also known
to an enlarged thyroid gland (goitre) or other as stress-activated protein kinase (SAPK),
iodine deficiency disorders including mental belongs to the family of MAP kinases.
retardation and stunted growth in babies and Jurkat cells A line of T lymphocyte cells that
children. Iodine is found in dairy products, are used to study acute T cell leukaemia.
seafood, kelp, seaweeds, eggs, some vegeta- KB cell A cell line derived from a human carci-
bles and iodised salt. noma of the nasopharynx, used as an assay for
IP See Intraperitoneal. antineoplastic (antitumour) agents.
IP3R3 (Inositol 1,4,5-triphosphate receptor Kainate receptors Or KARs, are non-NMDA
type 3) is an intracellular calcium release (N-methyl-D-aspartate) ionotropic recep-
channel that mediates calcium release from tors which respond to the neurotransmitter
the endoplasmic reticulum. glutamate.
Iron (Fe) Is essential to most life forms and to Kaliuresis The presence of excess potassium
normal human physiology. In humans, iron is in the urine.
an essential component of proteins involved Kallikreins Peptidases (enzymes that cleave
in oxygen transport and for haemoglobin. It is peptide bonds in proteins), a subgroup of the
318 Medical Glossary
serine protease family; they liberate kinins Kininogen Either of two plasma α2-globulins
from kininogens. Kallikreins are targets of that are kinin precursors.
active investigation by drug researchers as Ki-67 Human protein associated with cell
possible biomarkers for cancer. proliferation.
Kaposi sarcoma A cancerous tumour of the Knockout Gene knockout is a genetic
connective tissues caused by the human technique in which an organism is engi-
herpesvirus 8 and is often associated with neered to carry genes that have been made
AIDS. inoperative.
Kaposi sarcoma herpesvirus (KSHV) Also Konzo Is an epidemic paralytic disease occur-
known as human herpesvirus 8, is a gamma ring in outbreaks in remote rural areas of low-
2 herpesvirus or rhadinovirus. It plays an income African countries.
important role in the pathogenesis of Kaposi Kunitz protease inhibitors A type of protein
sarcoma (KS), multicentric Castleman dis- contained in legume seeds which functions as
ease (MCD) of the plasma cell type and pri- a protease inhibitor.
mary effusion lymphoma and occurs in HIV Kupffer cells Are resident macrophages of the
patients. liver and play an important role in its normal
Karyolysis Dissolution and disintegration of physiology and homeostasis as well as partici-
the nucleus when a cell dies. pating in the acute and chronic responses of
Karyorrhexis Destructive fragmentation of the liver to toxic compounds.
the nucleus of a dying cell whereby its chro- l-Dopa (L-3,4-Dihydroxyphenylalanine) is an
matin disintegrates into formless granules. amino acid that is formed in the liver and con-
Keloids Benign dermal tumours characterised verted into dopamine in the brain.
by fibroblastic proliferation and excessive Labour Process of childbirth involving muscu-
accumulation of collagen. lar contractions.
Keratin A sulphur-containing protein which is Lacrimation Secretion and discharge of tears.
a major component in skin, hair, nails, hooves, Lactagogue An agent that increases or stimu-
horns and teeth. lates milk flow or production. Also called a
Keratinocyte Is the major constituent of the galactagogue.
epidermis, constituting 95% of the cells found Lactate dehydrogenase (LDH) Enzyme that
there. catalyses the conversion of lactate to pyruvate.
Keratinophilic Having an affinity for keratin. Lactation Secretion and Production of milk.
Keratitis Inflammation of the cornea. Lactic acidosis Is a condition caused by the
Keratoconjunctivitis sicca Also called kera- build-up of lactic acid in the body. It leads to
titis sicca, xerophthalmia or dry eye syn- acidification of the blood (acidosis) and is con-
drome (DES), is an eye disease characterised sidered a distinct form of metabolic acidosis.
by a deficiency of aqueous tear film over the LAK cell A lymphokine-activated killer cell,
surface of the eye and in the lining of the i.e. a white blood cell that has been stimulated
lids. to kill tumour cells.
Keratolysis Softening and separation of the Lamella In cell biology, it refers to numerous
horny layer of the epidermis. plate or disc-like structures at both a tissue and
Keratolytic Pertaining to keratolysis. cellular level.
Keratomalacia An eye disorder that leads to a Laminin A glycoprotein component of con-
dry cornea. nective tissue basement membrane that pro-
Kidney stones (Calculi) are hardened mineral motes cell adhesion.
deposits that form in the kidney. Laparoscopic cholecystectomy Is a procedure
Kinin Is any of various structurally related in which the gallbladder is removed by laparo-
polypeptides, such as bradykinin, that act scopic techniques.
locally to induce vasodilation and contraction Laparotomy A surgical procedure involv-
of smooth muscle. ing an incision through the abdominal wall
Medical Glossary 319
to gain access into the abdominal cavity. adj. Leucocytopenia Abnormal decrease in the
laparotomised. number of leucocytes (white blood cells) in
Larvacidal An agent which kills insect or par- the blood.
asite larva. Leucocytosis Increase in white blood cell
Laryngitis Is an inflammation of the larynx. count above its normal range.
Laxation Bowel movement. Leucoderma A skin abnormality characterised
Laxatives Substances that are used to promote by white spots, bands and patches on the skin;
bowel movement. they can also be caused by fungus and tinea.
LC 50 Median lethal concentration; see LD 50. Also see Vitiligo.
LD 50 Median lethal dose—the dose required Leucomyelopathy Any diseases involving the
to kill half the members of a tested popu- white matter of the spinal cord.
lation. Also called LC 50 (median lethal Leucopenia A decrease in the number of circu-
concentration). lating white blood cells.
LDL See Low-density lipoprotein. Leucorrhoea Commonly known as whites,
LDL cholesterol See Low-density lipoprotein. refers to a whitish discharge from the female
LDL receptor (LDLr) A low-density lipopro- genitals.
tein receptor gene. Leukemia, leukaemia A cancer of the blood
Lectins Are sugar-binding proteins that are or bone marrow and is characterised by an
highly specific for their sugar moieties, which abnormal proliferation (production by multi-
agglutinate cells and/or precipitate glycocon- plication) of blood cells, usually white blood
jugates. They play a role in biological recogni- cells (leucocytes).
tion phenomena involving cells and proteins. Leukemogenic Relating to leukaemia, causing
Leiomyoma Benign smooth muscle neoplasm leukaemia.
that is very rarely (0.1%) premalignant. Leukoplakia Condition characterised by white
Leishmaniasis A disease caused by protozoan spots or patches on mucous membranes, espe-
parasites that belong to the genus Leishmania cially of the mouth and vulva.
and is transmitted by the bite of certain spe- Leukotriene A group of hormones that cause
cies of sand fly. the inflammatory symptoms of hay-fever and
Lenitive Palliative; easing pain or discomfort. asthma.
Lenticular opacity Also known as or related Leydig cells Also called interstitial cells of
to cataract. Leydig, are found adjacent to the seminiferous
Leprosy A chronic bacterial disease of the skin tubules in the testicle. They produce testoster-
and nerves in the hands and feet and, in some one in response to luteinising hormone.
cases, the lining of the nose. It is caused by the Levarterenol See Norepinephrine.
Mycobacterium leprae. Also called Hansen’s LexA repressor Or repressor LexA, is repres-
disease. sor enzyme that represses SOS response genes
Leptin Is a 16-kDa protein hormone with coding for DNA polymerases required for
important effects in regulating body weight, repairing DNA damage
metabolism and reproductive function. Libido Sexual urge.
Lequesne algofunctional index Is a wide- Lichen planus A chronic mucocutaneous
spread international instrument (ten-question disease that affects the skin, tongue and oral
survey) and recommended by the World mucosa.
Health Organization (WHO) for outcome Ligroin A volatile, inflammable fraction of
measurement in hip and knee diseases such as petroleum, obtained by distillation and used
osteoarthritis. as a solvent.
Leucocyte White blood corpuscles, colourless, Limbic system Complex set of brain struc-
without haemoglobin that helps to combat tures, including the hypothalamus, amyg-
infection. dala, hippocampus, anterior thalamic nuclei,
320 Medical Glossary
septum, limbic cortex and fornix that control Lovo cells Colon cancer cells.
various functions such as emotion, behaviour, Low-density lipoprotein (LDL) Is a type of
motivation, memory and olfaction. lipoprotein that transports cholesterol and tri-
Liniment Liquid preparation rubbed on skin, glycerides from the liver to peripheral tissues.
used to relieve muscular aches and pains. High levels of LDL cholesterol can signal
Linterised starch Starch that has undergone medical problems like cardiovascular disease,
prolonged acid treatment. and it is sometimes called ‘bad cholesterol’.
Lipodiatic Having lipid and lipoprotein lower- LRP1 Low-density lipoprotein receptor-related
ing property. protein-1, plays a role in intracellular signal-
Lipodystrophy A medical condition charac- ling functions as well as in lipid metabolism.
terised by abnormal or degenerative condi- LTB4 A type of leukotriene, a major metabolite
tions of the body’s adipose tissue. in neutrophil polymorphonuclear leucocytes.
Lipofuscin Finely granular yellow-brown pig- It stimulates polymorphonuclear cell function
ment granules composed of lipid-containing (degranulation, formation of oxygen-centred
residues of lysosomal digestion. free radicals, arachidonic acid release and
Lipogenesis Is the process by which acetyl- metabolism). It induces skin inflammation.
CoA is converted to fats; adj. lipogenic. Luciferase Is a generic name for enzymes com-
Lipolysis Is the breakdown of fat stored in fat monly used in nature for bioluminescence.
cells in the body. Lumbago Is the term used to describe general
Liposomes Artificially prepared vesicles made lower back pain.
of lipid bilayer. Lung abscess Necrosis of the pulmonary tissue
Lipotoxicity Refers to tissue diseases that may and formation of cavities containing necrotic
occur when fatty acids spill over in excess debris or fluid caused by microbial infections.
of the oxidative needs of those tissues and Lusitropic An agent that affects diastolic
enhances metabolic flux into harmful path- relaxation.
ways of nonoxidative metabolism. Lutein A carotenoid, occurs naturally as yel-
Lipotropic Refers to compounds that help low or orange pigment in some fruits and leafy
catalyse the breakdown of fat during vegetables. It is one of the two carotenoids
metabolism in the body, e.g. chlorine and contained within the retina of the eye. Within
lecithin. the central macula, zeaxanthin predominates,
Lipoxygenase A family of iron-containing whereas in the peripheral retina, lutein pre-
enzymes that catalyse the dioxygenation of dominates. Lutein is necessary for good vision
polyunsaturated fatty acids in lipids contain- and may also help prevent or slow down ath-
ing a cis,cis-1,4-pentadiene structure. erosclerosis, the thickening of arteries, which
Lithiasis Formation of urinary calculi (stones) is a major risk for cardiovascular disease.
in the renal system (kidneys, ureters, the uri- Luteinising hormone (LH) A hormone pro-
nary bladder, urethra) can be of any one of duced by the anterior pituitary gland. In
several compositions. females, it triggers ovulation. In males, it
Lithogenic Promoting the formation of calculi stimulates the production of testosterone to
(stones). aid sperm maturation.
Lithontripic Removes stones from the kidney, Luteolysis Is the structural and functional deg-
gallbladder. radation of the corpus luteum (CL) that occurs
Liver X receptors Nuclear hormones that at the end of the luteal phase of both the
function as central transcriptional regulators estrous and menstrual cycles in the absence of
for lipid homeostasis. pregnancy. adj. luteolytic.
Lochia Vaginal discharge containing blood, Luteotorpic Stimulating the formation of the
mucus and uterine tissues, during the postpar- corpus luteum.
tum period Lymphadenitis The inflammation or enlarge-
Lotion A liquid suspension or dispersion of ment of a lymph node casued by microbial
chemicals for external application to the body. infection.
Medical Glossary 321
cine zipper transcription factor protein that MMP Matrix metalloproteinases, a group of
plays a role in the development, survival and peptidases involved in the degradation of the
function of melanocytes and osteoclast. extracellular matrix (ECM).
Microsomal PGE2 synthase Is the enzyme Mnestic Pertaining to memory.
that catalyses the final step in prostaglandin Molecular docking Is a key tool in structural
E2 (PGE2) biosynthesis. molecular biology and computer-assisted drug
Microvasculature The finer vessels of the design.
body, as the arterioles, capillaries and venules. Molluscidal Destroying molluscs like snails.
Micturition Urination, act of urinating. Molt 4 cells MOLT4 cells are lymphoblast-
Migraine A neurological syndrome charac- like in morphology and are used for studies of
terised by altered bodily perceptions; severe, apoptosis, tumour cytotoxicity and tumorige-
painful headaches; and nausea. nicity, as well as for antitumour testing.
Mimosine Is an alkaloid, β-3-hydroxy-4 pyri- Molybdenum (Mo) Is an essential element that
done amino acid; it is a toxic nonprotein free forms part of several enzymes such as xanthine
amino acid and is an antinutrient. oxidase involved in the oxidation of xanthine
Mineral apposition rate MAR, rate of addi- to uric acid and use of iron. Molybdenum
tion of new layers of mineral on the trabecular concentrations also affect protein synthe-
surfaces of bones. sis, metabolism and growth. Dietary sources
Mineralcorticoids A group of steroid hor- include meat, green beans, eggs, sunflower
mones that are secreted by the adrenal cortex seeds, wheat flour, lentils and cereal grain.
and regulate the balance of water and electro- Monoamine oxidase A (MAOA) Is an iso-
lytes (sodium, potassium) in the body. zyme of monoamine oxidase. It preferentially
Miscarriage Spontaneous abortion. deaminates norepinephrine (noradrenaline),
Mitochondrial complex I The largest enzyme epinephrine (adrenaline), serotonin and
in the mitochondrial respiratory oxidative dopamine.
phosphorylation system. Monoaminergic Of or pertaining to neurons
Mitochondrial permeability transition that secrete monoamine neurotransmitters
(MPT) Is an increase in the permeability of (e.g. dopamine, serotonin).
the mitochondrial membranes to molecules of Monoclonal antibodies Are produced by fus-
less than 1500 Da in molecular weight. MPT ing single antibody-forming cells to tumour
is one of the major causes of cell death in a cells grown in culture.
variety of conditions. Monocyte Large white blood cell that ingests
Mitogen An agent that triggers mitosis and microbes, other cells and foreign matter.
elicits all the signals necessary to induce cell Monogalactosyl diglyceride Are the major
proliferation. lipid components of chloroplasts.
Mitogenic Able to induce mitosis or Monorrhagia Is heavy bleeding and that is
transformation. usually defined as periods lasting longer than
Mitogenicity Process of induction of mitosis. 7 days or excessive bleeding.
Mitomycin A chemotherapy drug that is given Morbidity A diseased state or symptom or can
as a treatment for several different types of refer either to the incidence rate or to the prev-
cancer, including breast|, stomach, oesopha- alence rate of a disease.
gus and bladder cancers. Morelloflavone A biflavonoid extracted from
Mitosis Cell division in which the nucleus Garcinia dulcis, has shown antioxidative,
divides into nuclei containing the same num- antiviral and anti-inflammatory properties.
ber of chromosomes. Morphine The major alkaloid of opium and a
Mitral valve prolapse The most common potent narcotic analgesic.
heart valve abnormality. Symptoms could MTTP Microsomal triglyceride transfer pro-
include palpitations, shortness of breath, tein that is required for the assembly and
cough, fatigue, dizziness or anxiety, migraine secretion of triglyceride-rich lipoproteins
headaches and chest discomfort. from both enterocytes and hepatocytes.
Medical Glossary 325
stimuli by sending nerve signals to the spinal Nuclear factor erythroid 2-related factor 2
cord and brain. (Nrf2) A transcription factor that plays a
Non-osteogenic Fibromata of bone, a benign major role in response to oxidative stress by
tumour of bone which shows no evidence of binding to antioxidant-responsive elements
ossification. that regulate many hepatic phase I and II
Non-alcoholic fatty liver disease One cause enzymes as well as hepatic efflux transporters.
of a fatty liver, occurring when fat is depos- Nucleosomes Fundamental repeating subunits
ited (steatosis) in the liver not due to excessive of all eukaryotic chromatin, consisting of a
alcohol use DNA chain coiled around a core of histones.
Nootropics Are substances which are claimed Nulliparous Term used to describe a woman
to boost human cognitive abilities (the func- who has never given birth.
tions and capacities of the brain). Also pop- Nyctalopia Night blindness, impaired vision
ularly referred to as ‘smart drugs’, ‘smart in dim light and in the dark, due to impaired
nutrients’, ‘cognitive enhancers’ and ‘brain function of certain specialised vision cells.
enhancers’. Nystagmus Fast, involuntary movements of
Noradrenalin See Norepinephrine. the eyes.
Norepinephrine A substance, both a hormone Nycturia Excessive urination at night; espe-
and neurotransmitter, secreted by the adrenal cially common in older men.
medulla and the nerve endings of the sympa- Obsessive–compulsive disorder (OCD) A
thetic nervous system to cause vasoconstric- common psychiatric disorder defined by the
tion and increases in heart rate, blood pressure presence of obsessive thoughts and repetitive
and the sugar level of the blood. Also called compulsive actions; self-grooming.
levarterenol, noradrenalin. Occludin A novel integral membrane protein
Normoglycaemic Having the normal amount localising at tight junctions. cf. tight junction.
of glucose in the blood. Occlusion Closure or blockage (as of a blood
Normotensive Having normal blood pressure. vessel).
Nosebo A harmless substance that when taken Occlusive peripheral arterial disease
by a patient is associated with unpleasant or (PAOD) Also known as peripheral vas-
harmful effects due to negative expectations cular disease (PVD), or peripheral arterial
or the psychological state of the person. disease (PAD), refers to the obstruction of
Nosocomial infections Infections which are a large arteries not within the coronary, aortic
result of treatment in a hospital or a healthcare arch vasculature, or the brain. PVD can result
service unit, but secondary to the patient’s from atherosclerosis, inflammatory processes
original condition. leading to stenosis, an embolism, or thrombus
NPC1L1 Niemann–Pick C1-like 1 gene that formation.
plays a major role in cholesterol homeosta- Oculomotor nerve The third of twelve paired
sis. It is critical for the uptake of cholesterol cranial nerves.
across the plasma membrane of the intestinal Odds ratio A statistical measure of effect size,
enterocyte. describing the strength of association or non-
Nrf2 Nuclear erythroid 2-related factor 2, independence between two binary data values.
a transcription factor that activates ARE- Odontalgia Toothache. adj. odontalgic.
containing genes. Odontopathy Any disease of the teeth.
Nrf2/ARE pathway Plays an important role Oedema See Edema.
in inducing phase II detoxifying enzymes and Oestrogen Female hormone produced by
antioxidant proteins and has been considered the ovaries that play an important role in the
a potential target for cancer chemoprevention estrous cycle in women.
because it eliminates harmful reactive oxygen Oestrogen receptor (ER) Is a protein found in
species or reactive intermediates generated high concentrations in the cytoplasm of breast,
from carcinogens. uterus, hypothalamus and anterior hypophysis
Medical Glossary 329
cells; ER levels are measured to determine a play a crucial role in heart and brain func-
breast CA’s potential for response to hormonal tion and in normal growth and development.
manipulation. Linoleic acid (LA) is the main omega-6 fatty
Oestrogen receptor positive (ER+) Means acid in foods, accounting for 85% to 90% of
that oestrogen is causing the tumour to grow the dietary omega-6 PUFA. Other omega-6
and that the breast cancer should respond well acids include gamma-linolenic acid or GLA,
to hormone suppression treatments. sometimes called gamoleic acid, eicosadienoic
Oestrogen receptor negative (ER-) Tumour is acid, arachidonic acid and docosadienoic acid.
not driven by oestrogen and needs another test Omega-9 fatty acids Are not essential poly-
to determine the most effective treatment. unsaturated fatty acids that have in common
Oestrus Sexual excitement or heat of female; a final carbon–carbon double bond in the n−9
or period of this characterised by changes in position. Some n−9s are common components
the sex organs. of animal fat and vegetable oil. Two n−9 fatty
Oligoarthritis An inflammation of two, three acids important in industry are:
or four joints. Oleic acid (18, 1, n−9), which is a main compo-
Oligoasthenoteratozoospermia A combina- nent of olive oil
tion of asthenozoospermia (reduced sperm Erucic acid (22, 1, n−9), which is found in rape-
motility) and oligozoospermia (low spermato- seed, wallflower seed and mustard seed
zoon count). Oncogenes Genes carried by tumour viruses
Oligonucleosome A series of nucleosomes. that are directly and solely responsible for the
Oligospermia or oligozoospermia Refers to neoplastic (tumorous) transformation of host
semen with a low concentration of sperm, cells.
commonly associated with male infertility. Oncosis Accidental cell death, also referred to
Oliguria Decreased production of urine. swelling necrosis.
Oligoanuria Insufficient urine volume to allow Ophthalmia Severe inflammation of the eye,
for administration of necessary fluids, etc. or the conjunctiva or deeper structures of the
Omega-3 fatty acids Are essential polyunsatu- eye. Also called ophthalmitis.
rated fatty acids that have in common a final Ophthalmia (sympathetic) Inflammation of
carbon–carbon double bond in the n−3 position. both eyes following trauma to one eye.
Dietary sources of omega-3 fatty acids include Ophthalmopathy An auntoimmune disease
fish oil and certain plant/nut oils. The three where the thyroid gland is overactive leading
most nutritionally important omega-3 fatty to ocular manifestations.
acids are alpha-linolenic acid, eicosapentaenoic Opiate Drug derived from the opium plant.
acid (EPA) and docosahexaenoic acid (DHA). Opioid receptors A group of G protein-
Research indicates that omega-3 fatty acids are coupled receptors located in the brain and
important in health promotion and disease and various organs that bind opiates or opioid
can help prevent a wide range of medical prob- substances.
lems, including cardiovascular disease, depres- Oppilation Obstruction particularly of the
sion, asthma and rheumatoid arthritis. lower intestines.
Omega-6 fatty acids Are essential polyun- Optic placode An ectodermal placode from
saturated fatty acids that have in common a which the lens of the embryonic eye develops;
final carbon–carbon double bond in the n−6 also called lens placode.
position. Omega-6 fatty acids are considered ORAC (oxygen radical absorbance capac-
essential fatty acids (EFAs) found in vegeta- ity) A method of measuring antioxidant
ble oils, nuts and seeds. They are essential to capacities in biological samples.
human health but cannot be made in the body. Oral submucous fibrosis A chronic debilitat-
Omega-6 fatty acids—found in vegetable oils, ing disease of the oral cavity characterised by
nuts and seeds—are a beneficial part of a heart- inflammation and progressive fibrosis of the
healthy eating. Omega-6 and omega-3 PUFA submucosa tissues.
330 Medical Glossary
Oral thrush An infection of yeast fungus, bone that is insufficient to keep up with the
Candida albicans, in the mucous membranes rate of bone destruction.
of the mouth. Osteoporosis A disease of bone that leads to an
Orchidectomy Surgery to remove one or both increased risk of fracture.
testicles. Osteoprotegerin Also called osteoclastogene-
Orchidectomised With the testis removed. sis inhibitory factor (OCIF), a cytokine, which
Orchitis An acute painful inflammatory reac- can inhibit the production of osteoclasts.
tion of the testis secondary to infection by dif- Osteosacrcoma A malignant bone tumour.
ferent bacteria and viruses. Also called osteogenic sarcoma.
Orexigenic Increasing or stimulating the Otalgia Earache, pain in the ear.
appetite. Otic placode A thickening of the ectoderm on
Orofacial dyskinesia Abnormal involuntary the outer surface of a developing embryo from
movements involving muscles of the face, which the ear develops.
mouth, tongue, eyes and, occasionally, the Otitis Inflammation of the inner or outer parts
neck—may be unilateral or bilateral and con- of the ear.
stant or intermittent. Otitis media Inflammation of the middle ear.
Oropharyngeal Relating to the oropharynx. Otorrhea Running drainage (discharge) exit-
Oropharynx Part of the pharynx between the ing the ear.
soft palate and the epiglottis. Otopathy Disease of the ear.
Osmophobia A fear, aversion or psychological Ovariectomised With one or two ovaries
hypersensitivity to odours. removed.
Ostalgia, ostealgia Pain in the bones. Also Ovariectomy Surgical removal of one or both
called osteodynia. ovaries.
Osteoarthritis Is the deterioration of the joints Oxidation The process of adding oxygen to a
that becomes more common with age. compound, dehydrogenation or increasing the
Osteoarthrosis Chronic noninflammatory electro-negative charge.
bone disease. Oxidoreductase activity Catalysis of an oxi-
Osteoblast A mononucleate cell that is respon- dation–reduction (redox) reaction, a revers-
sible for bone formation. ible chemical reaction. One substrate acts as
Osteoblastic Relating to osteoblasts. a hydrogen or electron donor and becomes
Osteocalcin A noncollagenous protein found oxidised, while the other acts as hydrogen or
in bone and dentin, also refers to as bone electron acceptor and becomes reduced.
gamma-carboxyglutamic acid-containing Oxygen radical absorbance capacity
protein. (ORAC) A method of measuring antioxi-
Osteoclasts A kind of bone cell that removes dant capacities in biological samples.
bone tissue by removing its mineralised Oxytocic adj. hastening or facilitating child-
matrix. birth, especially by stimulating contractions
Osteoclastogenesis The production of of the uterus.
osteoclasts. Oxytocin Is a mammalian hormone that also
Osteodynia Pain in the bone. acts as a neurotransmitter in the brain. It is best
Osteogenic Derived from or composed of any known for its roles in female reproduction. It
tissue concerned in bone growth or repair. is released in large amounts after distension of
Osteomalacia Refers to the softening of the the cervix and vagina during labour and after
bones due to defective bone mineralisation. stimulation of the nipples, facilitating birth
Osteomyelofibrosis A myeloproliferative dis- and breastfeeding, respectively*****.
order in which fibrosis and sclerosis finally Oxyuriasis Infestation by pinworms.
lead to bone marrow obliteration. Ozoena Discharge of the nostrils caused by
Osteopenia Reduction in bone mass, usually chronic inflammation of the nostrils.
caused by a lowered rate of formation of new p.o. Per os, oral administration.
Medical Glossary 331
Perniosis An abnormal reaction to cold that eral reference work for pharmaceutical drug
occurs most frequently in women, children specifications.
and the elderly. Also called chilblains. Pharyngitis, pharyngolaryngi-
Per os (P.O.) Oral administration. tis Inflammation of the pharynx and the
Peroxisome proliferator-activated recep- larynx.
tors (PPARs) A family of nuclear recep- Pharyngolaryngeal Pertaining to the pharynx
tors that are involved in lipid metabolism, and larynx.
differentiation, proliferation, cell death and Pharyngopathy Disease of the pharynx.
inflammation. Phase II drug metabolising enzymes Play
Peroxisome proliferator-activated receptor an important role in biotransformation of
alpha (PPAR-alpha) A nuclear receptor endogenous compounds and xenobiotics to
protein, transcription factor and a major regu- more easily excretable forms as well as in the
lator of lipid metabolism in the liver. metabolic inactivation of pharmacologically
Peroxisome proliferator-activated receptor active compounds. Phase II drug metabolising
gamma (PPAR-γ) A type II nuclear recep- enzymes are mainly transferases.
tor protein that regulates fatty acid storage and Phenolics Class of chemical compounds
glucose metabolism. consisting of a hydroxyl group (−OH)
Pertussis Whooping cough, sever cough. bonded directly to an aromatic hydrocarbon
Peyers patches Patches of lymphoid tissue or group.
lymphoid nodules on the walls of the ileal- Pheochromocytoma Is a rare neuroendo-
small intestine. crine tumour that usually originates from
PGE-2 Prostaglandin E2, a hormone-like sub- the adrenal glands’ chromaffin cells, causing
stance that is released by blood vessel walls in overproduction of catecholamines, powerful
response to infection or inflammation that acts hormones that induce high blood pressure and
on the brain to induce fever. other symptoms.
Phagocytes Are the white blood cells that pro- Phlebitis Is an inflammation of a vein, usually
tect the body by ingesting (phagocytosing) in the legs.
harmful foreign particles, bacteria and dead or Phlegm Abnormally viscid mucus secreted by
dying cells. adj. phagocytic. the mucosa of the respiratory passages during
Phagocytosis Is a process the human body uses certain infectious processes.
to destroy dead or foreign cells. Phlegmon A spreading, diffuse inflammation
Phantosmia A form of olfactory hallucination. of the soft or connective tissue due to infec-
Pharmacognosis The branch of pharmacology tion by streptococci bacteria.
that studies the composition, use and history Phonophobia Fear of loud sound.
of drugs. Phoroglucinol A white, crystalline compound
Pharmacodynamics Branch of pharmacol- used as an antispasmodic, analytical reagent
ogy dealing with the effects of drugs and the and decalcifier of bone specimens for micro-
mechanism of their action. scopic examination.
Pharmacokinetics Branch of pharmacology Phosphatidylglycerol Is a glycerophospholipid
concerned with the movement of drugs within found in pulmonary active surface lipoprotein
the body including processes of absorption, and consists of an L-glycerol 3-phosphate
distribution, metabolism and excretion in the backbone ester bonded to either saturated or
body. unsaturated fatty acids on carbons 1 and 2.
Pharmacopoeia Authoritative treatise contain- Phosphatidylinositol 3-kinases (PI 3-kinases
ing directions for the identification of drug or PI3Ks) A group of enzymes involved in
samples and the preparation of compound cellular functions such as cell growth, prolif-
medicines, published by the authority of a eration, differentiation, motility, survival and
government or a medical or pharmaceuti- intracellular trafficking, which in turn are
cal society and in a broader sense is a gen- involved in cancer.
334 Medical Glossary
Pomade A thick oily dressing. the colon, and thus improve host health. cf.
Porphyria A disorder wherein the body cannot probiotics.
convert naturally occurring compounds (pro- Pre-eclampsia Toxic condition of pregnancy
phyrins) into haem which contains iron. characterised by high blood pressure, abnor-
Porphyrin Any of a class of water-soluble, mal weight gain, proteinuria and oedema.
nitrogenous biological pigments, derivatives Prenidatory phase Preimplantation phase.
of which include the haemoproteins. Prepubertal Before puberty; pertaining to
Postherpetic neuralgia (PHN) is neuralgia the period of accelerated growth preceding
(pain in the nerves) caused by the varicella gonadal maturity.
herpes zoster virus. The pain may last for Pregnane X receptor (PXR, NR1I2) is a
more than a month or more after a shingles ligand-activated transcription factor that
infection occurred. plays a role not only in drug metabolism and
Postpartum depression Depression after transport but also in various other biological
pregnancy; also called postnatal depression. processes.
Postprandial After mealtime. Pregnenolone A steroid hormone produced by
Postural hypotension Also called orthostatic the adrenal glands, involved in the steroido-
hypotension—a condition of low blood pres- genesis of other steroid hormones like proges-
sure that can cause dizziness. terone, mineralocorticoids, glucocorticoids,
Potassium (K) Is an element that is essential androgens and oestrogens.
for the body’s growth and maintenance. It Prenidatory Referring to the time period
is necessary to keep a normal water balance between fertilisation and implantation.
between the cells and body fluids, for cellular Prenylated flavones Flavones with an isopre-
enzyme activities, and plays an essential role nyl group in the 8 position, has been reported
in the response of nerves to stimulation and to have good anti-inflammatory properties.
in the contraction of muscles. Potassium is Presyncopal sensation State consisting of
found in many plant foods and fish (tuna, hali- light-headedness, muscular weakness, blurred
but): chard, mushrooms, spinach, fennel, kale, vision and feeling faint.
mustard greens, Brussels sprouts, broccoli, Primiparous Relating to a woman who has
cauliflower, cabbage winter squash, eggplant, given birth once.
cantaloupe, tomatoes, parsley, cucumber, bell Proangiogenic Promote angiogenesis (forma-
pepper, turmeric, ginger root, apricots, straw- tion and development of new blood vessels).
berries, avocado and banana. Probiotication Enhancement with beneficial
Poultice Is a soft moist mass, often heated and probiotic bacteria such as Lactobacillus spe-
medicated, that is spread on cloth over the skin cies that can prevent the growth of intestinal
to treat an aching, inflamed or painful part of pathogenic microflora.
the body. Also called cataplasm. Probiotics Are dietary supplements and live
PPARs Peroxisome proliferator-activated microorganisms containing potentially benefi-
receptors—a group of nuclear receptor pro- cial bacteria or yeasts that are taken into the
teins that function as transcription factors alimentary system for healthy intestinal func-
regulating the expression of genes. tions. cf. prebiotics.
PR interval Is the time (in seconds) from Proctitis An inflammation of the rectum that
the beginning of the P wave (onset of atrial causes discomfort, bleeding and, occasionally,
depolarisation) to the beginning of the QRS a discharge of mucus or pus.
complex. Procyanidin Also known as proathocyanidin,
Prebiotics A category of functional food, oligomeric proathocyanidin, leukocyanidin
defined as non-digestible food ingredients and leucoanthocyanin, is a class of flavanols
that beneficially affect the host by selec- found in many plants. It has antioxidant activ-
tively stimulating the growth and/or activ- ity and plays a role in the stabilisation of col-
ity of one or a limited number of bacteria in lagen and maintenance of elastin.
Medical Glossary 337
Progestational Of or relating to the phase with muscular contraction and the inflamma-
of the menstrual cycle immediately follow- tion response such as swelling, pain, stiffness,
ing ovulation, characterised by secretion of redness and warmth.
progesterone. Prostaglandin E2 (PEG-2) One of the prosta-
Proglottid One of the segments of a tapeworm. glandins, a group of hormone-like substances
Prognosis Medical term to describe the likely that participate in a wide range of body func-
outcome of an illness. tions such as the contraction and relaxation of
Prokinetic Or gastroprokinetic, substance that smooth muscle, the dilation and constriction
enhances gastrointestinal motility by increas- of blood vessels, control of blood pressure and
ing the frequency of contractions in the small modulation of inflammation.
intestine or making them stronger. Prostaglandin E synthase An enzyme that
Prolactin A hormone produced by the pituitary in humans is encoded by the glutathione-
gland; it stimulates the breasts to produce milk dependent PTGES gene.
in pregnant women. It is also present in males, Prostanoids Term used to describe a subclass
but its role is not well understood. of eicosanoids (products of COX pathway)
Prolapse A common condition where the blad- consisting of the prostaglandins (mediators of
der, uterus and/or bowel protrudes into the inflammatory and anaphylactic reactions), the
vagina. thromboxanes (mediators of vasoconstriction)
Prolapsus To fall or slip out of place. and the prostacyclins (active in the resolution
Prolapus ani Eversion of the lower portion of phase of inflammation.)
the rectum and protruding through the anus, Prostanoid EP 4 A prostaglandin receptor that
common in infancy and old age. may be involved in the neonatal adaptation of
Proliferating cell nuclear antigen (PCNA) A circulatory system, osteoporosis as well as ini-
new marker to study human colonic cell tiation of skin immune responses.
proliferation. Prostate A gland that surrounds the urethra at
Proliferative vitreoretinopathy (PVR) Is a the bladder in the male.
blinding retinal condition. It involves the for- Prostate cancer A disease in which cancer
mation of pathological membranes, which dis- develops in the prostate, a gland in the male
lodges the retina and thereby compromises an reproductive system. Symptoms include pain,
individual’s ability to see. difficulty in urinating, erectile dysfunction
Prolyl-4-hydroxylase (P4H) Key enzyme in and other symptoms.
collagen synthesis. Prostate-specific antigen (PSA) A protein
Promastigote The flagellate stage in the devel- produced by the cells of the prostate gland.
opment of trypanosomatid protozoa, charac- Protein kinase C (PKC) A family of enzymes
terised by a free anterior flagellum. involved in controlling the function of other
Promyelocytic leukaemia A subtype of acute proteins through the phosphorylation of
myelogenous leukaemia (AML), a cancer of hydroxyl groups of serine and threonine
the blood and bone marrow. amino acid residues on these proteins. PKC
Pro-oxidants Chemicals that induce oxidative enzymes play important roles in several signal
stress, either through creating reactive oxygen transduction cascades.
species or inhibiting antioxidant systems. Protein tyrosine phosphatase (PTP) A group
Prophylaxis Prevention or protection against of enzymes that remove phosphate groups
disease. from phosphorylated tyrosine residues on
Proptosis See Exophthalmos. proteins.
Prostacyclin A prostaglandin that is a metabo- Proteinase A protease (enzyme) involved in
lite of arachidonic acid, inhibits platelet aggre- the hydrolytic breakdown of proteins, usually
gation and dilates blood vessels. by splitting them into polypeptide chains.
Prostaglandins A family of C 20 lipid com- Proteinuria Means the presence of an excess
pounds found in various tissues, associated of serum proteins in the urine.
338 Medical Glossary
Proteolysis Cleavage of the peptide bonds in Psychotropic Capable of affecting the mind,
protein forming smaller polypeptides. adj. emotions and behaviour.
proteolytic. PTEN Phosphatase and tensin homolog, a
Proteomics The large-scale study of proteins, tumour suppressor gene.
particularly their structures and functions. Ptosis Also known as drooping eyelid; caused
Prothrombin Blood-clotting protein that is by weakness of the eyelid muscle and damage
converted to the active form, factor IIa, or to the nerves that control the muscles or loose-
thrombin, by cleavage. ness of the skin of the upper eyelid.
Prothyroid Good for thyroid function. P13-K Is a lipid kinase enzyme involved in the
Protheolithic Proteolytic. See Proteolysis. regulation of a number of cellular functions
Proto-oncogene A normal gene which, when such as cell growth, proliferation, differentia-
altered by mutation, becomes an oncogene tion, motility, survival and intracellular traf-
that can contribute to cancer. ficking, which in turn are involved in cancer.
Prurigo A general term used to describe itchy P13-K/AKT signalling pathway Shown to be
eruptions of the skin. important for an extremely diverse array of
Pruritis Defined as an unpleasant sensation on the cellular activities—most notably cellular pro-
skin that provokes the desire to rub or scratch the liferation and survival.
area to obtain relief; itch, itching. adj. pruritic. Pthysis Silicosis with tuberculosis.
PSA Prostate-specific antigen, a protein which Ptosis Drooping of the upper eye lid.
is secreted into ejaculate fluid by the healthy PTP Protein tyrosine phosphatase.
prostate. One of its functions is to aid sperm PTPIB Protein tyrosine phosphatase 1B.
movement. P21 Also known as cyclin-dependent kinase
Pseudoaldosteronism Is a medical condition inhibitor 1 or CDK-interacting protein 1, is a
characterised by hypertension, reduced aldo- potent cyclin-dependent kinase inhibitor.
sterone secretion, hypokalemia and metabolic Puerperal Pertaining to childbirth.
acidosis and associated with low plasma renin Puerperium Postpartum period.
activity. Pulmonary embolism A blockage (blood clot)
Pseudohyperaldosteronism (also pseudoal- of the main artery of the lung.
dosteronism) Is a medical condition that Purgative A substance used to cleanse or
mimics hyperaldosteronism. Like hyperaldo- purge, especially causing the immediate evac-
steronism, it produces hypertension associated uation of the bowel.
with low plasma renin activity and metabolic Purpura Is the appearance of red or purple dis-
alkalosis associated with hypokalemia. colourations on the skin that do not blanch on
Pseudohypoaldosteronism A hereditary disor- applying pressure. Also called peliosis.
der of infancy characterised by severe salt and Purulent Containing pus discharge.
water depletion and other signs of aldosterone Purulent sputum Sputum containing, or con-
deficiency, although aldosterone secretion is sisting of, pus.
normal or increased; causes include aldoste- Pustule Small, inflamed, pus-filled lesions.
rone receptor defects and renal dysfunction. Pyelitis Acute inflammation of the pelvis of the
Psoriasis A common chronic, non-contagious kidney caused by bacterial infection.
autoimmune dermatosis that affects the skin Pyelonephritis An ascending urinary tract
and joints. infection that has reached the pyelum (pelvis)
Psychoactive Having effects on the mind or of the kidney.
behaviour. Pyoderma Bacterial skin infection.
Psychonautics Exploration of the psyche by Pyodermatitis Refers to inflammation of the
means of approaches such as meditation, skin.
prayer, lucid dreaming, brain wave entrain- Pyorrhea See Periodontitis.
ment, etc. Pyretic Referring to fever.
Psychotomimetic Hallucinogenic. Pyrexia Fever of unknown origin.
Medical Glossary 339
Pyridoxal A chemical form of vitamin B6. See Rash A temporary eruption on the skin; see
Vitamin B6. Uticaria.
Pyridoxamine A chemical form of vitamin Reactive oxygen species Species such as
B6. See Vitamin B6. superoxide, hydrogen peroxide and hydroxyl
Pyridoxine A chemical form of vitamin B6. radical. At low levels, these species may func-
See Vitamin B6. tion in cell signalling processes. At higher
Pyrolysis Decomposition or transformation of levels, these species may damage cellular
a compound caused by heat. adj. pyrolytic. macromolecules (such as DNA and RNA)
PYY peptide A 36-amino acid peptide and participate in apoptosis (programmed cell
secreted by L cells of the distal small intestine death).
and colon that inhibits gastric and pancreatic Rec A Is a 38-kDa Escherichia coli protein
secretion. essential for the repair and maintenance of
QSR complex Series of deflections in an elec- DNA.
trocardiogram that represent electrical activity Receptor for advanced glycation end prod-
generated by ventricular depolarisation prior ucts (RAGE) Is a member of the immu-
to contraction of the ventricle. noglobulin superfamily of cell surface
QT interval Is a measure of the time between molecules; mediates neurite outgrowth and
the start of the Q wave and the end of the cell migration upon stimulation with its
T wave in the heart’s electrical cycle. A ligand, amphoterin.
prolonged QT interval is a biomarker for Recticulocyte Non-nucleated stage in the
ventricular tachyarrhythmias and a risk factor development of the red blood cell.
for sudden death. Recticulocyte lysate Cell lysate produced
Quorum sensing (QS) The control of gene from reticulocytes, used as an in-vitro transla-
expression in response to cell density, is used tion system.
by both Gram-negative and Gram-positive Recticuloendothelial system Part of the
bacteria to regulate a variety of physiological immune system, consists of the phagocytic
functions. cells located in reticular connective tissue,
Radiculitis Inflammation of the radicle of a primarily monocytes and macrophages.
nerve. Recurrent aphthous stomatitis, or RAS Is a
Radiodermatitis A skin disease associated common, painful condition in which recurring
with prolonged exposure to ionising radiation. ovoid or round ulcers affect the oral mucosa.
Radiolysis The dissociation of molecules by Redox homeostasis Is considered as the cumu-
radiation. lative action of all free radical reactions and
Radioprotective Serving to protect or aiding antioxidant defences in different tissues.
in protecting against the injurious effect of Refrigerant A medicine or an application for
radiations. allaying heat, fever or its symptoms.
RAD23B UV excision repair protein RAD23 Renal calculi Kidney stones.
homolog B Renal interstitial fibrosis Damage sustained
RAGE Is the receptor for advanced glycation by the kidneys’ renal tubules and interstitial
end products, a multiligand receptor that prop- capillaries due to accumulation of extracellu-
agates cellular dysfunction in several inflam- lar waste in the wall of the small arteries and
matory disorders, in tumours and in diabetes. arterioles.
RANKL Receptor activator of nuclear factor Renal resistive index (RRI) Measures the
kappa-B ligand, a type II membrane protein resistance of renal arterial flow to the kidney.
and a member of the tumour necrosis factor Renin Also known as an angiotensinogenase,
(TNF) superfamily. is an enzyme that participates in the body’s
RAS See Renin–angiotensin system or recur- renin–angiotensin system (RAS). It regulates
rent aphthous stomatitis. the body’s mean arterial blood pressure.
340 Medical Glossary
Renin–angiotensin system (RAS) Also called and occurs commonly in children after a viral
the renin–angiotensin–aldosterone system infection.
(RAAS), is a hormone system that regulates Rhabdomyolysis Breakdown of muscle fibres
blood pressure and water (fluid) balance. leading to the release of muscle fibre content
Reperfusion The restoration of blood flow to (myoglobin) into the bloodstream.
an organ or tissue that has had its blood supply Rheumatic Pertaining to rheumatism or to
cut off, after a heart attack. abnormalities of the musculoskeletal system.
Reporter gene A transfected gene that pro- Rheumatism, rheumatic disorder, rheumatic
duces a signal, such as green fluorescence, diseases Refers to various painful medical
when it is expressed. conditions which affect bones, joints, muscles
Resistin A cysteine-rich protein secreted by and tendons. Rheumatic diseases are charac-
adipose tissue of mice and rats. terised by the signs of inflammation—redness,
Resolutive A substance that induces subsid- heat, swelling and pain.
ence of inflammation. Rheumatoid arthritis (RA) Is a chronic, sys-
Resolvent Reduce inflammation or swelling. temic autoimmune disorder that most com-
Respiratory burst Is the rapid release of monly causes inflammation and tissue damage
reactive oxygen species (superoxide radical in joints (arthritis) and tendon sheaths, together
and hydrogen peroxide) from different with anaemia.
cells. Rhinitis Irritation and inflammation of some
Resorb To absorb or assimilate a product of the internal areas of the nose and the primary
body such as an exudate or cellular growth. symptom of rhinitis is a runny nose.
Restenosis Is the reoccurrence of stenosis, Rhinopathy Disease or malformation of the
a narrowing of a blood vessel, leading to nose.
restricted blood flow. Rhinoplasty Is surgery to repair or reshape the
Resveratrol Is a phytoalexin produced natu- nose.
rally by several plants when under attack Rhinorrhea Commonly known as a runny
by pathogens such as bacteria or fungi. It is nose, characterised by an unusually significant
a potent antioxidant found in red grapes and amount of nasal discharge.
other plants. Rhinosinusitis Inflammation of the nasal cav-
Reticuloendothelial system Part of the ity and sinuses.
immune system that consists of the phagocytic Rho GTPases Rho-guanosine triphosphate
cells located in reticular connective tissue. hydrolase enzymes are molecular switches
Also called macrophage system or mononu- that regulate many essential cellular processes,
clear phagocyte system. including actin dynamics, gene transcription,
Retinal ischemia Is a common cause of visual cell cycle progression and cell adhesion.
impairment and blindness. Ribosome-inactivating proteins Proteins that
Retinitis pigmentosa (RP) An inherited, are capable of inactivating ribosomes.
degenerative eye disease that causes severe Rickets Is a softening of the bones in children
vision impairment and may lead to blindness. potentially leading to fractures and deformity.
Retinol A form of vitamin A; see Vitamin A. Ringworm Dermatophytosis, a skin infection
Retinoblastoma protein A tumour suppressor caused by fungus.
protein that is dysfunctional in several major Roborant Restoring strength or vigour, a tonic.
cancers. Rotavirus The most common cause of infec-
Retinopathy A general term that refers to tious diarrhoea (gastroenteritis) in young chil-
some form of noninflammatory damage to the dren and infants, one of several viruses that
retina of the eye. causes infections called stomach flu.
Revulsive Counterirritant, used for swellings. Rubefacient A substance for external appli-
Reye’s syndrome A potentially fatal disease cation that produces redness of the skin, e.g.
that has numerous detrimental effects to by causing dilation of the capillaries and an
many organs, especially the brain and liver, increase in blood.
Medical Glossary 341
Serotonin A monoamine neurotransmitter syn- Sialyation Reaction with sialic acid or its deriv-
thesised in serotonergic neurons in the central atives; used especially with oligosaccharides.
nervous system. Sialyltransferases Enzymes that transfer sialic
Sepsis Is a potentially fatal medical condition acid to nascent oligosaccharide.
characterised by a whole-body inflammatory Sickle cell disease Is an inherited blood dis-
response (called a systemic inflammatory order that affects red blood cells. People with
response syndrome or SIRS) that is triggered sickle cell disease have red blood cells that
by an infection. contain mostly haemoglobin S, an abnormal
Septicaemia A systemic disease associated type of haemoglobin. Sometimes these red
with the presence and persistence of patho- blood cells become sickle shaped (crescent
genic microorganisms or their toxins in the shaped) and have difficulty passing through
blood. small blood vessels.
Sequelae A pathological condition resulting Side stitch Is an intense stabbing pain under
from a prior disease, injury or attack. the lower edge of the ribcage that occurs while
Sexual potentiator Increases sexual activity exercising.
and potency and enhances sexual performance Signal transduction cascade Refers to a
due to increased blood flow and efficient series of sequential events that transfer a sig-
metabolism. nal through a series of intermediate molecules
Sexually transmitted diseases until final regulatory molecules, such as tran-
(STD) Infections that are transmitted scription factors, and is modified in response
through sexual activity. to the signal.
SGOT, serum glutamic oxaloacetic transami- Silicon (Si) Is required in minute amounts by
nase An enzyme that is normally present in the body and is important for the development
liver and heart cells. SGOT is released into of healthy hair and the prevention of nervous
blood when the liver or heart is damaged. Also disorders. Lettuce is the best natural source of
called aspartate transaminase (AST). silicon.
SGPT, serum glutamic pyruvic transami- Sinapism Signifies an external application, in
nase An enzyme normally present in serum the form of a soft plaster, or poultice.
and body tissues, especially in the liver; it is Sinusitis Inflammation of the nasal sinuses.
released into the serum as a result of tissue SIRC cells Statens Seruminstitut rabbit cornea
injury, also called alanine transaminase (ALT). (SIRC) cell line.
Shiga-like toxin A toxin produced by the bacte- SIRT 1 Stands for sirtuin (silent mating type
rium Escherichia coli which disrupts the func- information regulation 2 homolog) 1. It is an
tion of ribosomes, also known as verotoxin. enzyme that deacetylates proteins that contrib-
Shiga toxigenic Escherichia coli ute to cellular regulation.
(STEC) Comprises a diverse group of Sirtuin Also called Sir2 proteins, a class of
organisms capable of causing severe gastroin- proteins that possess either histone deacety-
testinal disease in humans. lase or mono-ribosyltransferase activity.
Shiga toxin A toxin produced by the bacterium 6-Keto-PGF1 alpha A physiologically active
Shigella dysenteriae, which disrupts the func- and stable hydrolysis product of epoprostenol,
tion of ribosomes. found in nearly all mammalian tissues.
Shingles Skin rash caused by the zoster virus Sjögren’s syndrome An autoimmune dis-
(same virus that causes chicken pox) and is ease that mainly affects the eyes and sali-
medically termed herpes zoster. vary glands, but can affect different parts of
Sialogogue Salivation promoter, a substance the body. Symptoms include dry and itchy
used to increase or promote the excretion of eyes, a dry mouth, thirst and swallowing
saliva. difficulties.
Sialoproteins Glycoproteins that contain sialic Skp1 (S-Phase kinase-associated protein 1) is a
acid as one of their carbohydrates. core component of SCF ubiquitin ligases and
Sialorrhea Excessive production of saliva. mediates protein degradation.
Medical Glossary 343
Superior mesenteric artery (SMA) Arises Synoviocyte Located in the synovial mem-
from the anterior surface of the abdominal brane; there are two types. Type A cells are
aorta, just inferior to the origin of the celiac more numerous, have phagocytic characteris-
trunk, and supplies the intestine from the tics and produce degradative enzymes. Type B
lower part of the duodenum to the left colic cells produce synovial fluid, which lubricates
flexure and the pancreas. the joint and nourishes the articular cartilage.
Superoxidae mutase (SOD) Antioxidant Syphilis Is perhaps the best known of all the
enzyme. STDs. Syphilis is transmitted by direct con-
Suppuration The formation of pus, the act of tact with infection sores, called chancres,
becoming converted into and discharging pus. syphilitic skin rashes or mucous patches on
Supraorbital Located above the orbit of the the tongue and mouth during kissing, necking,
eye. petting or sexual intercourse. It can also be
Sural nerve Sensory nerve comprising col- transmitted from a pregnant woman to a fetus
lateral branches off of the common tibial and after the fourth month of pregnancy.
common fibular nerve. System lupus erythematosus A long-term
SYK, spleen tyrosine kinase Is a human autoimmune disorder that may affect the
protein and gene. Syk plays a similar role in skin, joints, kidneys, brain and other organs.
transmitting signals from a variety of cell sur- Symptoms may include chest pain, fatigue,
face receptors including CD74, Fc receptor fever, hair loss, mailasie, mouth sores, sensi-
and integrins. tivity to sunlight, skin rash (butterfly rash).
Sympathetic nervous system The part of the Systolic The blood pressure when the heart is
autonomic nervous system originating in the contracting. It is specifically the maximum
thoracic and lumbar regions of the spinal cord arterial pressure during contraction of the left
that in general inhibits or opposes the physi- ventricle of the heart.
ological effects of the parasympathetic ner- T cells Or T lymphocytes, a type of white blood
vous system, as in tending to reduce digestive cell that plays a key role in the immune system
secretions or speed up the heart. and attacks virus-infected cells, foreign cells
Sympathomimetic Mimicking the effects of and cancer cells.
impulses conveyed by adrenergic postgangli- Tachyarrhythmia Any disturbance of the
onic fibres of the sympathetic nervous system. heart rhythm in which the heart rate is abnor-
Synaptic plasticity The ability of neurons mally increased.
to change the number and strength of their Tachycardia A false heart rate applied to
synapses. adults to rates over 100 beats per minute.
Synaptogenesis The formation of synapses. Tachykinins Neuropeptide transmitters that
Synaptoneurosomes Purified synapses con- are widely distributed and active in the central
taining the pre- and postsynaptic termini. nervous system and periphery, rapidly acting
Synaptosomes Isolated terminal of a neuron. secretagogues, and cause smooth muscle con-
Syncope Fainting, sudden loss of conscious- traction and vasodilation (hypotension).
ness followed by the return of wakefulness. Tachyphylaxia A decreased response to a
Syndactyly Webbed toes, a condition where medicine given over a period of time so that
two or more digits are fused together. larger doses are required to produce the same
Syneresis Expulsion of liquid from a gel, as response.
contraction of a blood clot and expulsion of Tachypnea Abnormally fast breathing.
liquid. Taenia A parasitic tapeworm or flatworm of
Syngeneic Genetically identical or closely the genus Taenia.
related, so as to allow tissue transplant; immu- Taeniacide An agent that kills tapeworms.
nologically compatible. Tardive dyskinesia A disorder characterised
Synovial Lubricating fluid secreted by syno- by repetitive, involuntary, purposeless move-
vial membranes, as those of the joints. ments in the body such as grimacing, tongue
346 Medical Glossary
protrusion, lip smacking, puckering and purs- Tetter Any of a number of skin diseases.
ing of the lips and rapid eye blinking. Rapid, TGF-beta Transforming growth factor beta is
involuntary movements of the limbs, torso and a protein that controls proliferation, cellular
fingers may also occur. differentiation and other functions in most
Tau Is a class of microtubule-associated pro- cells.
tein (MAP) in neuronal and glial cells. Th cells or T helper cells A subgroup of lym-
Tau-1 (Ser198/199/202), pS396 (Ser396) and phocytes that helps other white blood cells in
pS214 (Ser214) epitopes Serine phosphor- immunologic processes.
ylation sites of tau-1. Th 1 cells Helper cells that play an important
Tau phosphorylation Plays an important role role in the immune system.
in neurodegenerative diseases and regulated Th 17 cells A subset of T helper cells produc-
by protein kinases and phosphatases. ing interleukin 17.
TBARS See thiobarbituric acid reactive Thalassemia major Is a genetic blood disorder
substances. that causes the body to manufacture an abnor-
TCA cycle See Tricarboxylic acid cycle. mal form of haemoglobin.
TCID50 Median tissue culture infective dose; Thelarche The beginning of secondary (post-
that amount of a pathogenic agent that will natal) breast development, usually occurring
produce pathological change in 50% of cell at the beginning of puberty in girls.
cultures. Thermogenic Tending to produce heat, applied
Telencephalon The cerebral hemispheres, the to drugs or food (fat burning food).
largest divisions of the human brain. Thermogenesis Is the process of heat produc-
Teletherapy A noninvasive procedure using tion in organisms.
external beam radiotherapy treatments. Thermonociceptors Or thermal nociceptors,
Telomerase Enzyme that acts on parts of chro- sensory receptors that are stimulated by nox-
mosomes known as telomeres. ius heat or cold at various temperatures.
Temporomandibular joint disorder (TMJD or Thiobarbituric acid reactive substances
TMD syndrome) A disorder characterised (TBARS) A well-established method for
by acute or chronic inflammation of the tem- screening and monitoring lipid peroxidation.
poromandibular joint that connects the man- Thixotropy The property exhibited by certain
dible to the skull. gels of becoming fluid when stirred or shaken
Tendonitis Is inflammation of a tendon. and returning to the semisolid state upon
Tenesmus A strong desire to defaecate. standing.
Teras (Medicine) a grossly malformed and Thoracodynia Pain in the chest.
usually nonviable fetus. plural terata. 3-β-HSD (Or 3-β-hydroxysteroid
Teratogen Is an agent that can cause malforma- dehydrogenase/δ-5-4 isomerase) is an enzyme
tions of an embryo or fetus. adj. teratogenic. that catalyses the synthesis of progesterone
Testicular torsion Twisting of the spermatic from pregnenolone.
cord, which cuts off the blood supply to the 3-Nitrotyrosine (3-NT) protein Used as a
testicle and surrounding structures within the marker for oxidative damage or nitrosative
scrotum. stress.
Tetanus An acute, potentially fatal disease Thrombocythaemia A blood condition char-
caused by tetanus bacilli multiplying at the acterised by a high number of platelets in the
site of an injury and producing an exotoxin blood.
that reaches the central nervous system pro- Thrombocytopenia A condition when the
ducing prolonged contraction of skeletal mus- bone marrow does not produce enough plate-
cle fibres. Also called lockjaw. lets (thrombocytes) like in leukaemia.
Tete Acute dermatitis caused by both bacterial Thromboembolism Formation in a blood ves-
and fungal infections. sel of a clot (thrombus) that breaks loose and
Tetraparesis Weakness of muscles of all four is carried by the blood stream to plug another
limbs. vessel. cf. deep vein thrombosis.
Medical Glossary 347
Uterotrophic Causing an effect on the uterus. VCAM-1 (vascular cell adhesion mole-
Uterus Womb. cule-1) Also known as CD106, contains
Vaginal dystrophy A condition in which the six or seven immunoglobulin domains and
outer part of the vagina becomes dry and the is expressed on both large and small vessels
skin thickens or thins. only after the endothelial cells are stimulated
Vaginitis Infectious or non-infectious inflam- by cytokines.
mation of the vaginal mucosa. VEGF Vascular endothelial growth factor.
Vaginopathy Any disease of the vagina. Venereal disease (VD) Term given to the dis-
Vagotomy The surgical cutting of the vagus eases syphilis and gonorrhoea.
nerve to reduce acid secretion in the stomach. Venule A small vein, especially one joining
Vagus nerve A cranial nerve, that is, a nerve capillaries to larger veins.
connected to the brain. The vagus nerve has Vermifuge A substance used to expel worms
branches to most of the major organs in the from the intestines.
body, including the larynx, throat, windpipe, Verotoxin S Shiga-like toxin produced by
lungs, heart and most of the digestive system. Escherichia coli, which disrupts the function
Variola Or smallpox, a contagious disease of ribosomes, causing acute renal failure.
unique to humans, caused by either of two Verruca A contagious and painful wart on the
virus variants, Variola major and Variola sole of the foot.
minor. The disease is characterised by fever, Verruca plana Is a reddish-brown or flesh-
weakness and skin eruption with pustules that coloured, slightly raised, flat-surfaced, well-
form scabs that leave scars. demarcated papule on the hand and face, also
Varicose veins Are veins that have become called flat wart.
enlarged and twisted. Verruca vulgaris Small painless warts on the
Vasa vasorum Is a network of small blood ves- skin caused by the human papillomavirus.
sels that supply large blood vessels. pl. vasa Vertigo An illusory, sensory perception that
vasori. the surroundings or one’s own body is revolv-
Vascular cell adhesion molecule (VCAM) A ing; dizziness.
part of the immunoglobulin superfamily. Very low-density lipoprotein (VLDL) A type
They are important in inflammation, immune of lipoprotein made by the liver. VLDL is one
responses and intracellular signalling events. of the five major groups of lipoproteins (chy-
Vascular endothelial growth factor lomicrons, VLDL, intermediate-density lipo-
(VEGF) A polypeptide chemical produced protein, low-density lipoprotein, high-density
by cells that stimulates the growth of new lipoprotein (HDL)) that enable fats and cho-
blood vessels. lesterol to move within the water-based solu-
Vasculitis Group of disorders that destroy tion of the bloodstream. VLDL is converted
blood vessels by inflammation. in the bloodstream to low-density lipoprotein
Vasculogenesis The process of blood vessel (LDL).
formation occurring by a de novo production Vesical calculus Calculi (stones) in the urinary
of endothelial cells. bladder
Vasoconstrictor Drug that causes constriction Vesicant A substance that causes tissue
of blood vessels. blistering.
Vasodilator Drug that causes dilation or relax- Vestibular Relating to the sense of balance.
ation of blood vessels. Vestibular disorders Includes symptoms of
Vasodilatory Causing the widening of the dizziness, vertigo and imbalance; it can be
lumen of blood vessels. result from or worsened by genetic or envi-
Vasomotor symptoms Menopausal symptoms ronmental conditions.
characterised by hot flushes and night sweats. Vestibular schwannoma Also called acoustic
Vasospasm Refers to a condition in which neuroma is a benign tumour that may develop
blood vessels spasm, leading to vasoconstric- from an overproduction of Schwann cells that
tion and subsequently to tissue ischemia and press on the hearing and balance nerves in the
death (necrosis). inner ear.
Medical Glossary 351
Vestibular system Includes parts of the inner zyme in fatty acid metabolism. Deficiency
ear and brain that process sensory informa- causes paresthesia.
tion involved with controlling balance and eye Vitamin B6 water-soluble vitamin, exists in
movement. three major chemical forms: pyridoxine, pyri-
Vibrissa Stiff hairs that are located especially doxal and pyridoxamine. Vitamin B6 is needed
about the nostrils. in enzymes involved in protein metabolism,
Vimentin A type III intermediate filament pro- red blood cell metabolism, efficient func-
tein that is expressed in mesenchymal cells. tioning of nervous and immune systems and
Viremia A medical condition where viruses haemoglobin formation. Deficiency causes
enter the bloodstream and hence have access anaemia and peripheral neuropathy.
to the rest of the body. Vitamin B 7 Also called biotin or vitamin H,
Visceral fat Intra-abdominal fat, is located an essential water-soluble vitamin, is involved
inside the peritoneal cavity, packed in between in the synthesis of fatty acids, amino acids and
internal organs and torso. glucose, in energy metabolism. Biotin pro-
Visual entopia Visual disturbances motes normal health of sweat glands, bone
Vitamin Any complex, organic compound, marrow, male gonads, blood cells, nerve tis-
found in various food or sometimes synthe- sue, skin and hair. Deficiency causes dermati-
sised in the body, required in tiny amounts and tis and enteritis.
are essential for the regulation of metabolism, Vitamin B9 Also called folic acid, an essen-
normal growth and function of the body. tial water-soluble vitamin. Folate is especially
Vitamin A Retinol, fat-soluble vitamins that important during periods of rapid cell division
play an important role in vision, bone growth, and growth such as infancy and pregnancy.
reproduction, cell division and cell differentia- Deficiency during pregnancy is associated
tion, and help regulate the immune system in with birth defects such as neural tube defects.
preventing or fighting off infections. Vitamin Folate is also important for production of red
A that is found in colourful fruits and veg- blood cells and prevents anaemia. Folate is
etables is called provitamin A carotenoid. needed to make DNA and RNA, the building
They can be made into retinol in the body. blocks of cells. It also helps prevent changes
Deficiency of vitamin A results in night blind- to DNA that may lead to cancer.
ness and keratomalacia. Vitamin B12 A water-soluble vitamin, also
Vitamin B1 Also called thiamine, water-soluble called cobalamin as it contains the metal
vitamins, dissolve easily in water and, in gen- cobalt. It helps maintain healthy nerve cells
eral, are readily excreted from the body they and red blood cells and DNA production.
are not readily stored; consistent daily intake Vitamin B12 is bound to the protein in food.
is important. It functions as coenzyme in the Deficiency causes megaloblastic anaemia.
metabolism of carbohydrates and branched Vitamin C Also known as ascorbic acid is an
chain amino acids and other cellular processes. essential water-soluble vitamin. It functions as
Deficiency results in beriberi disease. cofactor for reactions requiring reduced cop-
Vitamin B2 Also called riboflavin, an essential per or iron metalloenzyme and as a protective
water-soluble vitamin that functions as coen- antioxidant. Deficiency of vitamin C causes
zyme in redox reactions. Deficiency causes scurvy.
ariboflavinosis. Vitamin D A group of fat-soluble, prohormone
Vitamin B3 Comprises niacin and niacina- vitamin, the two major forms of which are
mide, water-soluble vitamin that functions vitamin D2 (or ergocalciferol) and vitamin D3
as coenzyme or co-substrate for many redox (or cholecalciferol). Vitamin D obtained from
reactions and is required for energy metabo- sun exposure, food and supplements is biolog-
lism. Deficiency causes pellagra. ically inert and must undergo two hydroxyl-
Vitamin B5 Also called pantothenic acid, a ations in the body for activation. Vitamin D is
water-soluble vitamin that functions as coen- essential for promoting calcium absorption in
352 Medical Glossary
the gut and maintaining adequate serum cal- Vomitive Substance that causes vomiting.
cium and phosphate concentrations to enable Vulnerary (Wound healer) a substance used to
normal growth and mineralisation of bone heal wounds and promote tissue formation.
and prevent hypocalcaemic tetany. Deficiency Vulva-vaginal erythema Abnormal redness
causes rickets and osteomalacia. Vitamin D and inflammation of the skin in the vagina.
has other roles in human health, including Wart An infectious skin tumour caused by a
modulation of neuromuscular and immune viral infection.
function, reduction of inflammation and mod- Welt See Wheal.
ulation of many genes encoding proteins that Wheal A firm, elevated swelling of the skin.
regulate cell proliferation, differentiation and Also called a weal or welt.
apoptosis. White fat White adipose tissue (WAT) in mam-
Vitamin E Is the collective name for a group mals, store of energy. cf. brown fat.
of fat-soluble compounds and exists in eight Whitlow Painful infection of the hand involv-
chemical forms (alpha-, beta-, gamma- and ing one or more fingers that typically affects
delta-tocopherol and alpha-, beta-, gamma- the terminal phalanx.
and delta-tocotrienol). It has pronounced Whooping cough Acute infectious disease
antioxidant activities stopping the formation usually in children caused by a Bacillus bac-
of reactive oxygen species when fat under- terium and accompanied by catarrh of the
goes oxidation and helps prevent or delay the respiratory passages and repeated bouts of
chronic diseases associated with free radicals. coughing.
Besides its antioxidant activities, vitamin E is Wnt proteins Are a diverse family of secreted
involved in immune function, cell signalling, lipid-modified signalling glycoproteins that
regulation of gene expression and other meta- are 350–400 amino acids in length
bolic processes. Deficiency is very rare but Wnt signalling pathway Is a network of pro-
can cause mild haemolytic anaemia in new- teins involved in embryogenesis and cancer
born infants. and also in normal physiological processes.
Vitamin K A group of fat-soluble vitamin and X-Linked agammaglobulinemia Also known
consists of vitamin K1 which is also known as as X-linked hypogammaglobulinemia, XLA,
phylloquinone or phytomenadione (also called Bruton-type agammaglobulinemia, Bruton
phytonadione) and vitamin K2 (menaquinone, syndrome or sex-linked agammaglobulin-
menatetrenone). Vitamin K plays an important emia; a rare x-linked genetic disorder that
role in blood clotting. Deficiency is very rare affects the body’s ability to fight infection.
but can cause bleeding diathesis. Xanthine oxidase A flavoprotein enzyme con-
Vitamin P A substance or mixture of taining a molybdenum cofactor (Moco) and
substances obtained from various plant (Fe2S2) clusters, involved in purine metabo-
sources, identified as citrin or a mixture of lism. In humans, inhibition of xanthine oxi-
bioflavonoids, thought to but not proven dase reduces the production of uric acid and
to be useful in reducing the extent of prevents hyperuricaemia and gout.
haemorrhage. Xanthones Unique class of biologically
Vitiligo A chronic skin disease that causes loss active phenol compounds with the molecular
of pigment, resulting in irregular pale patches formula C13H8O2 possessing antioxidant
of skin. It occurs when the melanocytes, cells properties, discovered in the mangosteen
responsible for skin pigmentation, die or are fruit.
unable to function. Also called leucoderma. Xenobiotics A chemical (as a drug, pesti-
Vitreoretinopathy See Proliferative cide or carcinogen) that is foreign to a living
vitreoretinopathy. organism.
VLA-4 Very late antigen-4, expressed by most Xenograft A surgical graft of tissue from one
leucocytes, but it is observed on neutrophils species to an unlike species.
under special conditions. Xerophthalmia A medical condition in which
VLDL See Very low-density lipoproteins. the eye fails to produce tears.
Medical Glossary 353
Xerostomia Dryness in the mouth due to lack for good vision and is one of the two carot-
of saliva production. enoids contained within the retina of the eye.
Yaws An infectious tropical infection of the Within the central macula, zeaxanthin pre-
skin, bones and joints caused by the spirochete dominates, whereas in the peripheral retina,
bacterium Treponema pertenue, characterised lutein predominates.
by papules and papilloma with subsequent Zinc (Zn) Is an essential mineral for health.
deformation of the skins, bone and joints; also It is involved in numerous aspects of cellular
called framboesia. metabolism: catalytic activity of enzymes,
yGCN5 A histone acetyl transferase (HAT) immune function, protein synthesis, wound
that plays a role in regulation of transition, cell healing, DNA synthesis and cell division. It
cycle progression and differentiation. also supports normal growth and development
Yellow fever Is a viral disease that is transmit- during pregnancy, childhood and adolescence
ted to humans through the bite of infected and is required for proper sense of taste and
mosquitoes. Illness ranges in severity from an smell. Dietary sources include beans, nuts,
influenza-like syndrome to severe hepatitis and pumpkin seeds, sunflower seeds, whole wheat
haemorrhagic fever. Yellow fever virus (YFV) bread and animal sources.
is maintained in nature by mosquito-borne ZK1 Kruppel-type zinc finger protein—binds
transmission between nonhuman primates. DNA and, through this binding, regulates
Zeaxanthin A common carotenoid, found nat- gene transcription.
urally as coloured pigments in many fruit veg- ZO1 protein A high molecular weight tight
etables and leafy vegetables. It is important junction-associated protein.
Scientific Glossary
Abaxial Facing away from the axis, as of the leaves, buds arising elsewhere than in the axils
surface of an organ. of leaves.
Abortive Imperfectly formed. Adventive Not native to and not fully estab-
Abscission Shedding of leaves, flowers or lished in a new habitat or environment; locally
fruits following the formation of the abscis- or temporarily naturalised, e.g. an adventive
sion zone. weed.
Acaulescent Lacking a stem, or stem very Aestivation Refers to positional arrangement
much reduced. of the floral parts in the bud before it opens.
Accrescent Increasing in size after flowering Akinete A thick-walled dormant cell derived
or with age. from the enlargement of a vegetative cell. It
Achene A dry, small, one-seeded, indehiscent serves as a survival structure.
fruit formed from a superior ovary of one car- Alfisols Soil with a clay-enriched subsoil
pel as in sunflower. and relatively high native fertility, having
Acid soil Soil that maintains a pH of less than undergone only moderate leaching, con-
7.0. taining aluminium, iron and with at least
Acidulous Acid or sour in taste. 35% base saturation, meaning that calcium,
Actinomorphic Having radial symmetry, capa- magnesium and potassium are relatively
ble of being divided into symmetrical halves abundant.
by any plane, referring to a flower, calyx or Alkaline soil Soil that maintains a pH above
corolla. 7.0, usually containing large amounts of cal-
Aculeate Having sharp prickles. cium, sodium and magnesium, and is less
Acuminate Tapering gradually to a sharp point. soluble than acidic soils.
Acute (Botany) tapering at an angle of less Alkaloids Naturally occurring bitter, com-
than 90° before terminating in a point as of plex organic-chemical compounds containing
leaf apex and base. basic nitrogen and oxygen atoms and having
Adaxial Side closest to the stem axis. various pharmacological effects on humans
Adelphous Having stamens united together by and other animals.
their filaments. Alternate Leaves or buds that are spaced along
Adherent Touching without organic fusion as opposite sides of stem at different levels.
of floral parts of different whorls. Allomorphic With a shape or form different
Adnate United with another unlike part as of from the typical.
stamens attached to petals. Alluvial soil A fine-grained fertile soil depos-
Adpressed Lying close to another organ but ited by water flowing over flood plains or in
not fused to it. river beds.
Adventitious Arising in abnormal positions, Alluvium Soil or sediments deposited by a
e.g. roots arising from the stem, branches or river or other running water.
Amplexicaul Clasping the stem as base of cer- Apically Towards the apex or tip of a structure.
tain leaves. Apiculate Ending abruptly in a short, sharp,
Anatomising Interconnecting network as small point.
applied to leaf veins. Apiculum A short, pointed, flexible tip.
Andisols Are soils formed in volcanic ash Apocarpous Carpels separate in single indi-
and containing high proportions of glass and vidual pistils.
amorphous colloidal materials. Apopetalous With separate petals, not united
Androdioecious With male flowers and bisex- to other petals.
ual flowers on separate plants. Aposepalous With separate sepals, not united
Androecium Male parts of a flower; compris- to other sepals.
ing the stamens of one flower. Appendiculate Having small appendages
Androgynophore A stalk bearing both the Appressed Pressed closely to another structure
androecium and gynoecium above the peri- but not fused or united.
anth of the flower. Aquatic A plant living in or on water for all or
Androgynous With male and female flowers in a considerable part of its life span.
distinct parts of the same inflorescence. Arachnoid (Botany) formed of or covered
Andromonoecious Having male flowers and with long, delicate hairs or fibres.
bisexual flowers on the same plant. Arborescent Resembling a tree; applied to
Angiosperm A division of seed plants with the nonwoody plants attaining tree height and to
ovules borne in an ovary. shrubs tending to become tree-like in size.
Annual A plant which completes its life cycle Arbuscular mycorrhiza (AM) A type of
within a year. mycorrhiza in which the fungus (of the phy-
Annular Shaped like or forming a ring. lum Glomeromycota) penetrates the cortical
Annulus Circle or ringlike structure or mark- cells of the roots of a vascular plant and forms
ing; the portion of the corolla which forms a unique structures such as arbuscules and vesi-
fleshy, raised ring. cles. These fungi help plants to capture nutri-
Anthelate An open, paniculate cyme. ents such as phosphorus and micronutrients
Anther The part of the stamen containing pol- from the soil.
len sac which produces the pollen. Archegonium A flask-shaped female repro-
Antheriferous Containing anthers. ductive organ in mosses, ferns and other
Anthesis The period between the opening of related plants.
the bud and the onset of flower withering. Areolate With areolae.
Anthocarp A false fruit consisting of the true Areole (Botany) a small, specialised, cushion-
fruit and the base of the perianth. like area on a cactus from which hairs, glo-
Anthocyanidins Are common plant pig- chids, spines, branches or flowers may arise;
ments. They are the sugar-free counterparts of an irregular angular specs marked out on a
anthocyanins. surface, e.g. fruit surface. pl. areolae.
Anthocyanins A subgroup of antioxidant fla- Aril Specialised outgrowth from the funiculus
vonoids, are glucosides of anthocyanidins. (attachment point of the seed) (or hilum) that
They occur as water-soluble vacuolar pig- encloses or is attached to the seed. adj. arillate.
ments that may appear red, purple or blue Arillode A false aril; an aril originating from
according to pH in plants. the micropyle instead of from the funicle or
Antipetala Situated opposite petals. chalaza of the ovule, e.g. mace of nutmeg.
Antisepala Situated opposite sepals. Aristate Bristlelike part or appendage, e.g.
Antrorse Directed forward and upward. awns of grains and grasses.
Apetalous Lacking petals as of flowers with no Aristulate Having a small, stiff, bristlelike part
corolla. or appendage; a diminutive of aristate
Apical meristem Active growing point. A zone Articulate Jointed; usually breaking eas-
of cell division at the tip of the stem or the ily at the nodes or point of articulation into
root. segments.
356 Scientific Glossary
Ascending Arched upwards in the lower part Bifid Forked, divided into two parts.
and becoming erect in the upper part. Bifoliolate Having two leaflets.
Ascospore Spore produced in the ascus in Bilabiate Having two lips as of a corolla or
ascomycete fungi. calyx with segments fused into an upper and
Ascus Is the sexual spore-bearing cell pro- lower lip.
duced in ascomycete fungi. pl. asci. Bipinnate Twice pinnate; the primary leaflets
Asperulous Refers to a rough surface with being again divided into secondary leaflets.
short, hard projections. Bipinnatisect Refers to a pinnately compound
Attenuate Tapered or tapering gradually to a leaf, in which each leaflet is again divided into
point. pinnae.
Auricle An ear-like appendage that occurs at Biserrate Doubly serrate; with smaller, regu-
the base of some leaves or corolla. lar, asymmetric teeth on the margins of larger
Auriculate Having auricles. teeth.
Awn A hair-like or bristlelike appendage on a Bisexual Having both sexes, as in a flower
larger structure. bearing both stamens and pistil, hermaphro-
Axil Upper angle between a lateral organ, such dite or perfect.
as a leaf petiole, and the stem that bears it. Biternate Twice ternate; with three pinnae
Axile Situated along the central axis of an each divided into three pinnules.
ovary having two or more locules, as in axile Blade Lamina; part of the leaf above the sheath
placentation. or petiole.
Axillary Arising or growing in an axil. Blotched See Variegated.
Baccate Beery-like, pulpy or fleshy. Bole Main trunk of tree from the base to the
Barbate Bearded, having tufts of hairs. first branch.
Barbellae Short, stiff, hair-like bristles. adj. Brachyblast A short, axillary, densely crowded
barbellate. branchlet or shoot of limited growth, in which
Bark Is the outermost layers of stems and roots the internodes elongate little or not at all.
of woody plants. Bracket fungus Shelf fungus.
Basal Relating to, situated at, arising from or Bract A leaflike structure, different in form
forming the base. from the foliage leaves, associated with an
Basaltic soil Soil derived from basalt, a com- inflorescence or flower. adj. bracteate.
mon extrusive volcanic rock. Bracteate Possessing bracts.
Basidiospore A reproductive spore produced Bracteolate Having bracteoles.
by basidiomycete fungi. Bracteole A small, secondary, bract-like struc-
Basidium A microscopic, spore-producing ture borne singly or in a pair on the pedicel or
structure found on the hymenophore of fruit- calyx of a flower. adj. bracteolate.
ing bodies of basidiomycete fungi. Bran Hard outer layer of grain and comprises
Basifixed Attached by the base, as certain the aleurone and pericarp. It contains impor-
anthers are to their filaments. tant antioxidant, vitamins and fibre.
Basionym The synonym of a scientific name Bristle A stiff hair.
that supplies the epithet for the correct name. Bulb A modified underground axis that is short
Beak A prominent apical projection, especially and crowned by a mass of usually fleshy,
of a carpel or fruit. adj. beaked. imbricate scales. adj. bulbous.
Bearded Having a tuft of hairs. Bulbil A small bulb or bulb-shaped body, espe-
Berry A fleshy or pulpy indehiscent fruit from cially one borne in the leaf axil or an inflorescence
a single ovary with the seed(s) embedded in and usually produced for asexual reproduction.
the fleshy tissue of the pericarp. Bullate Puckered, blistered.
Biconvex Convex on both sides. Burr Type of seed or fruit with short, stiff
Biennial Completing the full cycle from ger- bristles or hooks or may refer to a deformed
mination to fruiting in more than one, but not type of wood in which the grain has been
more than two years. misformed.
Scientific Glossary 357
Bush Low, dense shrub without a pronounced Capsule A dry, dehiscent fruit formed from
trunk. two or more united carpels and dehiscing at
Buttress Supporting, projecting out- maturity by sections called valves to release
growth from base of a tree trunk as in some the seeds. adj. capsular.
Rhizophoraceae and Moraceae. Carinate Keeled.
Caducous Shedding or falling early before Carpel A simple pistil consisting of ovary,
maturity refers to sepals and petals. ovules, style and stigma. adj. carpellary.
Caespitose Growing densely in tufts or clumps; Carpogonium Female reproductive organ in
having short, closely packed stems. red algae. pl. carpogonia.
Calcareous Composed of or containing lime or Carpophore Part of the receptacle which is
limestone. lengthened between the carpels as a central
Calcrete A hardpan consisting gravel and sand axis; any fruiting body or fruiting structure of
cemented by calcium. a fungus.
Callus A condition of thickened raised mass of Cartilaginous Sinewy, having a firm, tough,
hardened tissue on leaves or other plant parts flexible texture (in respect of leaf margins).
often formed after an injury but sometimes a Caruncle (Bot) fleshy structure attached to the
normal feature. A callus also can refer to an seed of certain plants.
undifferentiated plant cell mass grown on a Caryopsis A simple dry, indehiscent fruit
culture medium. n. callosity. pl. calli, callosi- formed from a single ovary with the seed coat
ties. adj. callose. united with the ovary wall as in grasses and
Calyptra The protective cap or hood cov- cereals.
ering the spore case of a moss or related Cataphyll A reduced or scarcely developed
plant. leaf at the start of a plant’s life (i.e. cotyledons)
Calyptrate Operculate, having a calyptra. or in the early stages of leaf development.
Calyx Outer floral whorl usually consisting Catkin A slim, cylindrical, pendulous flower
of free sepals or fused sepals (calyx tube) and spike usually with unisexual flowers.
calyx lobes. It encloses the flower while it is Caudate Having a narrow, tail-like appendage.
still a bud. adj. calycine. Caudex Thickened, usually underground base
Calyx lobe One of the free upper parts of the of the stem.
calyx which may be present when the lower Caulescent Having a well-developed aerial
part is united into a tube. stem.
Calyx tube The tubular fused part of the calyx, Cauliflory Botanical term referring to plants
often cup shaped or bell shaped, when it is which flower and fruit from their main stems
free from the corolla. or woody trunks. adj. cauliflorus.
Campanulate Shaped like a bell refers to calyx Cauline Borne on the aerial part of a stem.
or corolla. Chaffy Having thin, membranous scales in the
Canaliculate Having groove or grooves. inflorescence as in the flower heads of the sun-
Candelabriform Having the shape of a tall flower family.
branched candle stick. Chalaza The basal region of the ovule where
Canescent Covered with short, fine whitish or the stalk is attached.
greyish hairs or down. Chamaephyte A low-growing perennial plant
Canopy Uppermost leafy stratum of a tree. whose dormant overwintering buds are borne
Cap See Pileus. at or just above the surface of the ground.
Capitate Growing together in a head. Also Chartaceous Papery, of paper-like texture.
means enlarged and globular at the tip. Chasmogamous Describing flowers in which
Capitulum A flower head or inflorescence hav- pollination takes place while the flower is
ing a dense cluster of sessile, or almost sessile, open.
flowers or florets. Chatoyant Having a velvety sheen or lustre.
358 Scientific Glossary
Cymbiform Boat shaped and elongated and Dichasium A cymose inflorescence in which
having the upper surface decidedly concave. the branches are opposite and approximately
Cyme An inflorescence in which the lateral equal. pl. dichasia. adj. dichasial.
axis grows more strongly than the main axis Dichotomous Divided into two parts.
with the oldest flower in the centre or at the Dicotyledon Angiosperm with two cotyledons.
ends. adj. cymose Didymous Arranged or occurring in pairs as of
Cymule A small cyme or one or a few flowers. anthers, having two lobes.
Cystidium A relatively large cell found on the Digitate Having digits or fingerlike projections.
hymenium of a basidiomycete, for example, Dikaryophyses Or dendrophydia, irregularly,
on the surface of a mushroom. strongly branched terminal hyphae in the
Cystocarp Fruitlike structure (sporocarp) hymenomycetes (class of basidiomycetes)
developed after fertilisation in the red algae. fungi.
Deciduous Falling off or shedding at maturity Dimorphic Having or occurring in two forms,
or a specific season or stage of growth. as of stamens of two different lengths or a
Decorticate To remove the bark, rind or husk plant, having two kinds of leaves.
from an organ; to strip of its bark; to come off Dioecious With male and female unisexual
as a skin. flowers on separate plants. cf. monoecious.
Decompound As of a compound leaf; consist- Diploid a condition in which the chromosomes
ing of divisions that are themselves compound. in the nucleus of a cell exist as pairs, one set
Decumbent Prostrate, laying or growing on the being derived from the female parent and the
ground but with ascending tips. cf. ascending, other from the male.
procumbent. Diplobiontic life cycle Life cycle that exhib-
Decurrent Having the leaf base tapering down its alternation of generations, which features
to a narrow wing that extends to the stem. spore-producing multicellular sporophytes
Decussate Having paired organs with succes- and gamete-producing multicellular gameto-
sive pairs at right angles to give four rows as of phytes. Mitoses occur in both the diploid and
leaves. haploid phases.
Deflexed Bent downwards. Diplontic life cycle Or gametic meiosis,
Degumming Removal of gum deposits (phos- wherein instead of immediately dividing mei-
phatides, entrained oil and meal particles) otically to produce haploid cells, the zygote
from crude edible oils traditionally done divides mitotically to produce a multicellular
with water. Water degumming process also diploid individual or a group of more diploid
removes hydrophilic substances such as sug- cells.
ars from the oil. Diplochory Seed dispersal involving two or
Dehisce To split open at maturity, as in a more modes.
capsule. Dipterocarpous Trees of the family
Dehiscent Splitting open at maturity to release Dipterocarpaceae, with two-winged fruit
the contents. cf. indehiscent. found mainly in tropical lowland rainforest.
Deltate Triangular shape. Disc (Botany) refers to the usually disc-
Deltoid Shaped like an equilateral triangle. shaped receptacle of the flower head in
Dendritic Branching from a main stem or axis Asteraceae; also the fleshy nectariferous
like the branches of a tree. organ usually between the stamens and
Dentate With sharp, rather coarse teeth per- ovary; also used for the enlarged style end
pendicular to the margin. in Proteaceae.
Denticulate Finely toothed. Disc floret The central, tubular four- or
Diageotropic The tendency of growing parts, five-toothed or lobed floret on the disc
such as roots, to grow at right angle to the line of an inflorescence, as of flower head of
of gravity. Asteraceae.
Diadelphous Having stamens in two bundles Disciform Flat and rounded in shaped. cf. dis-
as in Papilionaceae flowers. coid, radiate.
Scientific Glossary 361
Discoid Resembling a disc; having a flat, cir- Embryo (Botany) a minute rudimentary plant
cular form; disc shaped. cf. disciform, radiate. contained within a seed or an archegonium,
Discolorous Having two colours, as of a leaf composed of the embryonic axis (shoot end
which has different colours on the two sur- and root end).
faces. cf. concolorous. Endemic Prevalent in or peculiar to a particu-
Disomic Having one or more chromosomes lar geographical locality or region.
present twice but without the entire genome Endocarp The hard innermost layer of the
doubled. pericarp of many fruits.
Dispersal Dissemination of seeds. Endosperm Tissue that surrounds and nour-
Distal Site of any structure farthest from the ishes the embryo in the angiosperm seed. It
point of attachment. cf. proximal. contains starchy carbohydrates, proteins and
Distichous Referring to two rows of upright small amounts of vitamins and minerals.
leaves in the same plane. Endospermous Refers to seeds having an
Dithecous Having two thecae. endosperm.
Divaricate Diverging at a wide angle. Ensiform Shaped like the blade of a sword,
Domatium A part of a plant (e.g. a leaf) that long and narrow with sharp edges and a
has been modified to provide protection for pointed tip.
other organisms. pl. domatia. Endotrophic As of mycorrhiza obtaining
Dormancy A resting period in the life of a nutrients from inside.
plant during which growth slows or appears Ensilage The process of preserving green food
to stop. for livestock in an undried condition in airtight
Dorsal Referring to the back surface. conditions. Also called silaging.
Dorsifixed Attached to the back as of anthers. Entire Having a smooth, continuous margin
Drupaceous Resembling a drupe. without any incisions or teeth as of a leaf.
Drupe A fleshy fruit with a single seed enclosed Entisols Soils that do not show any profile
in a hard shell (endocarp) which is tissue development other than an A horizon.
embedded in succulent tissue (mesocarp) sur- Ephemeral Transitory, short lived.
rounded by a thin outer skin (epicarp). adj. Epicalyx A whorl of bracts, subtending and
drupaceous. resembling a calyx.
Drupelet A small drupe. Epicarp Outermost layer of the pericarp of a fruit.
Ebracteate Without bracts. Epicormic Attached to the corm.
Echinate Bearing stiff, stout, bristly, prickly Epicotyl The upper portion of the embryonic
hairs. axis, above the cotyledons and below the first
Edaphic Refers to plant communities that are true leaves.
distinguished by soil conditions rather than by Epigeal Above grounds with cotyledons raised
the climate. above ground.
Eglandular Without glands. cf. glandular. Epiparasite An organism parasitic on another
Elaioplasts A type of leucoplast that is special- that parasitizes a third.
ised for the storage of lipids in plants. Epipetalous Borne on the petals, as of stamens.
Elaiosome Fleshy lipid-rich structures that are Epiphyte A plant growing on, but not para-
attached to the seeds of many plant species. sitic on, another plant, deriving its moisture
Ellipsoid A three-dimensional shape; elliptic and nutrients from the air and rain, e.g. some
in outline. Orchidaceae. adj. epiphytic.
Elliptic Having a two-dimensional shape of an Epithet Name.
ellipse or flattened circle. Equitant In a loose fan pattern.
Elongate Extended, stretched out. Erect Upright, vertical.
Emarginate Refers to leaf with a broad, shal- Essential oils Volatile products obtained from
low notch at the apex. cf. retuse. a natural source; refers to volatile products
362 Scientific Glossary
obtained by steam or water distillation in a Floret One of the small individual flowers of
strict sense. sunflower family or the reduced flower of the
Etiolation To cause (a plant) to develop with- grasses, including the lemma and palea.
out chlorophyll by preventing exposure to Flower The sexual reproductive organ of flow-
sunlight. ering plants, typically consisting of gynoe-
Eutrophic Having waters rich in mineral and cium, androecium and perianth or calyx and/
organic nutrients that promote a proliferation or corolla and the axis bearing these parts.
of plant life, especially algae, which reduces Fluted As of a trunk with grooves and folds.
the dissolved oxygen content and often causes Fodder Plant material, fresh or dried fed to
the extinction of other organisms. animals.
Excentric Off the true centre. Foliaceous Leaflike.
Excrescence Abnormal outgrowth. Foliage Leaves of the plant.
Excurrent Projecting beyond the tip, as the Foliar Pertaining to a leaf.
midrib of a leaf or bract. Foliolate Pertaining to leaflets, used with a
Exserted Sticking out, protruding beyond number prefix to denote the number of leaflets.
some enclosing organ, as of stamens which Foliose Leaflike.
project beyond the corolla or perianth. Follicle (Botany) a dry fruit, derived from
Exstipulate Without stipules. cf. stipulate. a single carpel and dehiscing along one
Extra-floral Outside the flower. suture.
Extrose Turned outwards or away from the Forb Any herb that is not grass or grass-like.
axis as of anthers. cf. introrse, latrorse. Foveolate Surface pitted with shallow
Falcate Sickle shaped, crescent shaped. depressions.
Fascicle A cluster or bundle of stems, flowers Free central placentation The arrangement
and stamens. adj. fasciculate. of ovules on a central column that is not con-
Fasciclode Staminode bundles. nected to the ovary wall by partitions, as in the
Fastigiate A tree in which the branches grow ovaries of the carnation and primrose.
almost vertically. Frond The leaf of a fern or cycad.
Ferrosols Soils with an iron oxide content of Fruit Ripened ovary with adnate parts.
greater than 5%. Frutescent Shrubby.
Ferruginous Rust coloured, reddish-brown. Fugacious Shedding off early.
Fertile Having functional sexual parts which Fulvous Yellow, tawny.
are capable of fertilisation and seed produc- Funiculus (Botany) short stalk which attaches
tion. cf. sterile. the ovule to the ovary wall.
Filament The stalk of a stamen supporting and Fuscescent Dusky.
subtending the anther. Fusiform A three-dimensional shape; spindle
Filiform Having the form of or resembling a shaped, i.e. broad in the centre, but tapering at
thread or filament. both thick ends.
Fimbriate Fringed. Galea A part of the calyx or corolla having the
Fixed oils Non-volatile oils, triglycerides of form of a helmet.
fatty acids. Gall flower Short-styled flower that does not
Flaccid Limp and weak. develop into a fruit but is adapted for the
Flag leaf The uppermost leaf on the stem. development of a specific wasp within the
Flaky In the shape of flakes or scales. fruit, e.g. in the fig.
Flexuous Zigzagging, sinuous, bending, as of Gamete A reproductive cell that fuses with
a stem. another gamete to form a zygote. Gametes are
Floccose Covered with tufts of soft woolly hairs. haploid (they contain half the normal (diploid)
Floral tube A flower tube usually formed by number of chromosomes); thus when two
the basal fusion of the perianth and stamens. fuse, the diploid number is restored.
Scientific Glossary 363
Gametophyte The gamete-producing phase Groats Hulled, whole grains of various cere-
in a plant characterised by alternation of als, such as oats, wheat, barley or buckwheat;
generations. it includes the cereal germ, fibre-rich bran por-
Gamosepalous With sepals united or partially tion and endosperm of the grain.
united. Guttation The appearance of drops of xylem
Genome Complete set of genetic material of an sap on the tips or edges of leaves of some vas-
organism. cular plants, such as grasses and bamboos.
Geniculate Bent like a knee, refer to awns and Guttule Small droplet.
filaments. Gymnosperm A group of spermatophyte
Geocarpic Where the fruit is pushed into the seed-bearing plants with ovules on scales,
soil by the gynophore and matures. which are usually arranged in cone-like
Geophyte A plant that stores food in an under- structures and not borne in an ovary. cf.
ground storage organ, e.g. a tuber, bulb or rhi- angiosperm.
zome, and has subterranean buds which form Gynoecium The female organ of a flower;
aerial growth. a collective term for the pistil, carpel or
Geotextile Are permeable fabrics which, when carpels.
used in association with soil, have the ability Gynomonoecious Having female flowers
to separate, filter, reinforce, protect or drain. and bisexual flowers on the same plant. cf.
Germ Of cereal is the embryo of the seed or andromonoecious.
kernel. It contains vitamins B and E, folic Gynophore Stalk that bears the pistil/carpel.
acid, some protein, minerals and polyunsatu- Habit The general growth form of a plant,
rated fats. comprising its size, shape, texture and stem
Glabrescent Becoming glabrous. orientation, the locality in which the plant
Glabrous Smooth, hairless without grows..
pubescence. Halophyte A plant adapted to living in highly
Gland A secretory organ, e.g. a nectary, extra- saline habitats. Also a plant that accumulates
floral nectary or a gland tipped, hair-like or high concentrations of salt in its tissues. adj.
wartlike organ. adj. glandular. cf. eglandular. halophytic.
Glaucous Pale blue-green in colour, covered Hapaxanthic Refer to palms which flower
with a whitish bloom that rubs off readily. only once and then die. c.f. pleonanthic.
Gley soils A hydric soil which exhibits a Haploid Condition where nucleus or cell has a
greenish-blue-grey soil colour due to wetland single set of unpaired chromosomes; the hap-
conditions. loid number is designated as n.
Globose Spherical in shape. Haplontic life cycle Or zygotic meiosis
Globular A three-dimensional shape; spherical wherein meiosis of a zygote, immediately
or orbicular; circular in outline. after karyogamy, produces haploid cells
Glochids Tiny, finely barbed hair-like spines which produces more or larger haploid cells
found on the areoles of some cacti and other ending its diploid phase.
plants. Hastate Having the shape of an arrowhead but
Glochidiate Having glochids. with the basal lobes pointing outwards at right
Glochidote Plant having glochids. angles as of a leaf.
Glume One of the two small, sterile bracts at Hastula A piece of plant material at the junc-
the base of the grass spikelet, called the lower tion of the petiole and the leaf blade; the has-
and upper glumes, due to their position on the tula can be found on the top of the leaf, adaxial
rachilla. Also used in Apiaceae, Cyperaceae or the bottom, abaxial or both sides.
for the very small bracts on the spikelet in Heartwood Wood from the inner portion of a
which each flower is subtended by one floral tree.
glume. adj. glumaceous. Heliophilous Sun loving, tolerates high level
Grits Consist of coarsely ground corn, or of sunlight.
sometimes alkali-treated corn. Heliotropic Growing towards sunlight.
364 Scientific Glossary
Herb A plant which is nonwoody or woody at Homogamous Bearing flowers or florets that
the base only, the above-ground stems usually do not differ sexually. cf. heterogamous.
being ephemeral. adj. herbaceous. Homogeneous endosperm Endosperm with
Herbaceous Resembling a herb, having a habit even surface that lacks invaginations or infold-
of a herb. ings of the surrounding tissue.
Hermaphrodite Bisexual, bearing flowers Homomorphous Uniform, with only one
with both androecium and gynoecium in the form. cf. heteromorphous.
same flower. adj. hermaphroditic. Homosporous Producing one kind of
Heterocyst A differentiated cyanobacterial cell spores. Refer to the ferns and fern allies. cf.
that carries out nitrogen fixation. heterosporous.
Heterogamous Bearing separate male and Hormogonium A part of a filament of a cya-
female flowers, or bisexual and female flow- nobacterium that detaches and grows by cell
ers, or florets in an inflorescence or flower division into a new filament. pl. hormogonia.
head, e.g. some Asteraceae in which the ray Hurd fibre Long pith fibre of the stem.
florets may be neuter or unisexual and the disc Hyaline Colourless, almost transparent.
florets may be bisexual. cf. homogamous. Hybrid The first-generation progeny of the
Heteromorphous Having two or more distinct sexual union of plants belonging to different
forms. cf. homomorphous. taxa.
Heterophyllous Having leaves of different Hybridisation The crossing of individuals
forms. from different species or taxa.
Heterosporous Producing spores of two sizes, Hydathode A type of secretory tissue in leaves,
the larger giving rise to megagametophytes usually of Angiosperms, that secretes water
(female) and the smaller giving rise to micro- through pores in the epidermis or margin of
gametophytes (male). Refer to the ferns and the leaf.
fern allies. cf. homosporous. Hydrophilous Water loving; requiring water
Heterostylous Having styles of two different in order to be fertilised, referring to many
lengths or forms. aquatic plants.
Heterostyly The condition in which flowers on Hygrochastic Applied to plants in which the
polymorphous plants have styles of different opening of the fruits is caused by the absorp-
lengths, thereby facilitating cross-pollination. tion of water.
Hilar Of or relating to a hilum. Hygrophilous Living in water or moist places.
Hilum The scar on a seed, indicating the point Hymenial cystidia The cells of the hymenium
of attachment to the funiculus. develop into basidia or asci, while in others
Hirsute Bearing long coarse hairs. some cells develop into sterile cells called
Hispid Bearing stiff, short, rough hairs or cystidia.
bristles. Hymenium Spore-bearing layer of cells in cer-
Hispidulous Minutely hispid. tain fungi containing asci (ascomycetes) or
Histosol Soil comprising primarily of organic basidia (basidiomycetes).
materials, having 40 cm or more of organic Hypanthium Cup-like receptacles of some
soil material in the upper 80 cm. dicotyledonous flowers formed by the fusion
Hoary Covered with a greyish layer of very of the calyx, corolla and androecium that sur-
short, closely interwoven hairs. rounds the ovary which bears the sepals, petals
Holdfast An organ or structure of attachment, and stamens. adj. relating to or of the nature of
especially the basal, root-like formation by a hypanthium.
which certain seaweeds or other algae are Hypha Is a long, branching filamentous cell of
attached to a substrate. a fungus and also of unrelated Actinobacteria.
Holocarpic Having the entire thallus devel- pl. hyphae.
oped into a fruiting body or sporangium. Hypocotyl The portion of the stem below the
Homochromous Having all the florets of cotyledons.
the same colour in the same flower head cf. Hypodermis The cell layer beneath the epider-
heterochromous. mis of the pericarp.
Scientific Glossary 365
Hypogeal Below ground as of germination of Integuments Two distinct tissue layers that
seed. surround the nucellus of the ovule, forming
Hysteresis Refers to systems that may exhibit the testa or seed coat when mature.
path dependence. Intercalary Of growth, between the apex and
Imbricate Closely packed and overlapping. cf. the base; of cells, spores, etc., between two
valvate. cells.
Imparipinnate Pinnately compound with a Interfoliar Inter-leaf.
single terminal leaflet and hence with an odd Internode Portion of the stem, culm, branch
number of leaflets. cf. paripinnate. or rhizome between two nodes or points of
Inceptisols Old soils that have no accumulation attachment of the leaves.
of clays, iron, aluminium or organic matter. Interpetiolar As of stipules positioned
Incised Cut jaggedly with very deep teeth. between petioles of opposite leaves.
Included Referring to stamens which do not Intrastaminal Within the stamens.
project beyond the corolla or to valves which Intricate Entangled, complex.
do not extend beyond the rim of a capsular Introduced Not indigenous; not native to the
fruit. cf. exserted. area in which it now occurs.
Incurved Curved inwards; curved towards the Introrse Turned inwards or towards the axis or
base or apex. pistil as of anthers. cf. extrorse, latrorse.
Indefinite Numerous and variable in number. Involucre A whorl of bracts or leaves that
Indehiscent Not opening or splitting to surround one to many flowers or an entire
release the contents at maturity as of fruit. cf. inflorescence.
dehiscent. Involute Having the margins rolled inwards,
Indumentum covering of fine hairs or bristles referring to a leaf or other flat organ.
commonly found on external parts of plants. Jugate Of a pinnate leaf; having leaflets in
Indurate To become hard, often the hardening pairs.
developed only at maturity. Juvenile Young or immature, used here for leaves
Indusium An enclosing membrane, covering formed on a young plant which are different in
the sorus of a fern. Also used for the modified morphology from those formed on an older plant.
style end or pollen-cup of some Goodeniaceae Keel A longitudinal ridge, at the back of the leaf.
(including Brunoniaceae). adj. indusiate. Also the two lower fused petals of a ‘pea’ flower
Inferior Said of an ovary or fruit that has in the Papilionaceae, which form a boat-like
sepals, petals and stamens above the ovary. cf. structure around the stamens and styles, also
superior. called carina. adj. keeled. cf. standard, wing.
Inflated Enlarged and hollow except in the Labellum The modified lowest of the three
case of a fruit which may contain a seed. cf. petals forming the corolla of an orchid, usu-
swollen. ally larger than the other two petals, and often
Inflexed Bent or curved inwards or down- spurred.
wards, as petals or sepals. Lacerate Irregularly cleft.
Inflorescence A flower cluster or the arrange- Laciniate Fringed; having a fringe of slender,
ment of flowers in relation to the axis and to narrow, pointed lobes cut into narrow lobes.
each other on a plant. Lamella A gill-shaped structure: fine sheets of
Infrafoliar Located below the leaves. material held adjacent to one another.
Infraspecific Referring to any taxon below the Lamina The blade of the leaf or frond.
species rank. Lanate Woolly, covered with long hairs which
Infructescence The fruiting stage of an are loosely curled together like wool.
inflorescence. Lanceolate Lance shaped in outline, tapering
Infundibulum Funnel-shaped cavity or from a broad base to the apex.
structure. Landrace: landrace Plants adapted to the nat-
Inrolled Curved inwards. ural environment in which they grow, devel-
366 Scientific Glossary
oping naturally with minimal assistance or Limb The expanded portion of the calyx tube
guidance from humans and usually possess or the corolla tube, or the large branch of a
more diverse phenotypes and genotypes. They tree.
have not been improved by formal breeding Linear A two-dimensional shape, narrow with
programmes. nearly parallel sides.
Laterite Reddish-coloured soils rich in iron Linguiform Tongue shaped. cf. ligulate.
oxide, formed by weathering of rocks under Lipotubuloids Are cytoplasmic domains con-
oxidising and leaching conditions, commonly taining aggregates of lipid bodies surrounded
found in tropical and subtropical regions. adj. by a network of microtubules, which join one
lateritic. lipid body with the others.
Latex A milky, clear or sometimes coloured Lithosol A kind of shallow soils lacking well-
sap of diverse composition exuded by some defined horizons and composed of imperfectly
plants. weathered fragments of rock.
Latrorse Turned sideways, i.e. not towards or Littoral Of or on a shore, especially seashore.
away from the axis as of anthers dehiscing lon- Loam A type of soil made up of sand, silt and
gitudinally on the side. cf. extrorse, introrse. clay in relative concentration of 40–40–20%
Lax Loose or limp, not densely arranged or respectively.
crowded. Lobed Divided but not to the base.
Leaflet One of the ultimate segments of a com- Loculicidal Opening into the cells, when a ripe
pound leaf. capsule splits along the back.
Lectotype A specimen chosen after the origi- Loculus Cavity or chamber of an ovary. pl.
nal description to be the type. loculi.
Lemma The lower of two bracts (scales) of a Lodicules Two small structures below the
grass floret, usually enclosing the palea, lodi- ovary which, at flowering, swell up and force
cules, stamens and ovary. open the enclosing bracts, exposing the sta-
Lenticel Is a lens shaped opening that allows mens and carpel.
gases to be exchanged between air and the Lorate Strap shaped with obtuse tip.
inner tissues of a plant, commonly found on Lyrate Pinnately lobed, with a large terminal
young bark, or the surface of the fruit. lobe and smaller lateral ones which become
Lenticellate Dotted with lenticels. progressively smaller towards the base.
Lenticular Shaped like a biconvex lens. cf. Macronutrients Chemical elements which
lentiform. are needed in large quantities for growth and
Lentiform Shaped like a biconvex lens, cf. development by plants and include nitrogen,
lenticular. phosphorus, potassium and magnesium.
Leptomorphic Temperate, running bam- Maculate Spotted.
boo rhizome; usually thinner than the culms Mallee A growth habit in which several to
they support and the internodes are long and many woody stems arise separately from a
hollow. lignotuber; usually applied to certain low-
Liane A woody climbing or twining plant. growing species of Eucalyptus.
Ligneous Woody. Mangrove A distinctive vegetation type of
Lignotuber A woody, usually underground, trees and shrubs with modified roots, often
tuberous rootstock often giving rise to numer- viviparous, occupying the saline coastal
ous aerial stems. habitats that are subject to periodic tidal
Ligulate Small and tongue shaped or with a inundation.
little tongue-shaped appendage or ligule, star Marcescent Withering or to decay without
shaped as of florets of Asteraceae. falling off.
Ligule A strap-shaped corolla in the flowers Margin The edge of the leaf blade.
of Asteraceae; also a thin membranous out- Medulla The pith in the stems or roots of cer-
growth from the inner junction of the grass tain plants; or the central portion of a thallus
leaf sheath and blade. cf. ligulate. in certain lichens.
Scientific Glossary 367
Megasporangium The sporangium contain- Mollisols Soils with deep, high organic mat-
ing megaspores in fern and fern allies. cf. ter, nutrient-enriched surface soil (a horizon),
microsporangium. typically between 60 and 80 cm thick.
Megaspore The large spore which may develop Monadelphous Applied to stamens united by
into the female gametophyte in heterosporous their filaments into a single bundle.
ferns and fern allies. cf. microspore. Monocarpic Refer to plants that flower, set
Megasporophyll A leaflike structure that bears seeds and then die.
megasporangia. Monochasial A cyme having a single flower on
Megastrobilus Female cone, seed cone or each axis.
ovulate cone and contains ovules within Monocotyledon Angiosperm having one
which, when fertilised by pollen, become cotyledon.
seeds. The female cone structure varies Monoecious Having both male and female uni-
more markedly between the different conifer sexual flowers on the same individual plant.
families. cf. dioecious.
Meiosis The process of cell division that results Monoembryonic seed The seed contains only
in the formation of haploid cells from diploid one embryo, a true sexual (zygotic) embryo.
cells to produce gametes. Polyembryonic seed.
Mericarp A one-seeded portion of an initially Monolete A spore that has a simple linear scar.
syncarpous fruit (schizocarp) which splits Monopodial With a main terminal growing
apart at maturity. cf. coccus. point producing many lateral branches pro-
Meristem The region of active cell division gressively. cf. sympodial.
in plants, from which permanent tissue is Monostichous Forming one row.
derived. adj. meristematic Monotypic Of a genus with one species or a
-merous Used with a number prefix to denote family with one genus; in general, applied to
the basic number of the three outer floral any taxon with only one immediately subor-
whorls, e.g. a five-merous flower may have 5 dinate taxon.
sepals, 10 petals and 15 stamens. Montane Refers to highland areas located
Mesic Moderately wet. below the subalpine zone.
Mesocarp The middle layer of the fruit wall Mucilage A soft, moist, viscous, sticky secre-
derived from the middle layer of the carpel tion. adj. mucilaginous.
wall. cf. endocarp, exocarp and pericarp. Mucous (Botany) slimy.
Mesophytes Terrestrial plants which are Mucro A sharp, pointed part or organ, espe-
adapted to neither a particularly dry nor par- cially a sharp terminal point, as of a leaf.
ticularly wet environment. Mucronate Ending with a short, sharp tip or
Micropyle The small opening in a plant ovule mucro, resembling a spine. cf. cuspidate,
through which the pollen tube passes in order muticous.
to effect fertilisation. Mucronulate With a very small mucro; a
Microsporangium The sporangium con- diminutive of mucronate.
taining microspores in pteridophytes. cf. Mulch Protective cover of plant (organic) or
megasporangium. non-plant material placed over the soil, pri-
Microspore A small spore which gives rise marily to modify and improve the effects of
to the male gametophyte in heterosporous the local microclimate and to control weeds.
pteridophytes. Also for a pollen grain. cf. Multiple fruit A fruit that is formed from a
megaspore. cluster of flowers.
Midvein The main vascular supply of a Muricate Covered with numerous short hard
simple leaf blade or lamina. Also called outgrowths. cf. papillose.
midrib. Muriculate With numerous minute hard out-
Mitosis Is a process of cell division which growths; a diminutive of muricate.
results in the production of two daughter cells Muticous Blunt, lacking a sharp point. cf.
from a single parent cell. mucronate.
368 Scientific Glossary
MYB proteins Are a superfamily of transcrip- to which it was applied already has a name, or
tion factors that play regulatory roles in devel- because the name has already been applied to
opmental processes and defence responses in another plant.
plants. Nom. invalid. Nomen invalidum (Latin)
Mycorrhiza The mutualistic symbiosis (non- invalid name according to International Code
pathogenic association) between soilborne of Botanical Nomenclature.
fungi with the roots of higher plants. Nom. nud. Nomen nudum (Latin) the name
Mycorrhiza (vesicular arbuscu- of a taxon which has never been validated by
lar) Endomycorrhiza living in the roots of a description.
higher plants producing inter- and intracellu- Nom. rej. Nomen rejiciendum (Latin) name
lar fungal growth in root cortex and forming rejected in International Code of Botanical
specific fungal structures, referred to as vesi- Nomenclature.
cles and arbuscules. abbrev. VAM. Notho- (Subsp. or var.) prefix to the rank of a
Myrmecochory Seed dispersal by ants. hybrid taxon below the rank of species.
Native A plant indigenous to the locality or Nucellus Central portion of an ovule in which
region. the embryo sac develops.
Naviculate Boat shaped. Nucellar embryony A form of seed repro-
Necrotic Applied to dead tissue. duction in which the nucellar tissue which
Nectariferous Having one or more nectaries. surrounds the embryo sac can produce addi-
Nectary A nectar secretory gland; commonly tional embryos (polyembryony) which are
in a flower, sometimes on leaves, fronds or genetically identical to the parent plant. This
stems. is found in many citrus species and in mango.
Nervation Venation, a pattern of veins or Nut A dry indehiscent one-celled fruit with a
nerves as of leaf. hard pericarp.
Nixtamalisation Refers to a process for the Nutlet A small, one-seeded, indehiscent lobe
preparation of maize (corn), or other grains, in of a divided fruit.
which the grains are soaked and cooked in an Ob- Prefix meaning inversely or opposite to.
alkaline solution, usually limewater, and hulled. Obconic A three-dimensional shape; inversely
Node The joint between segments of a culm, conic; cone shaped, conic with the vertex
stem, branch or rhizome; the point of the stem pointing downwards.
that gives rise to the leaf and bud. Obcordate Inversely cordate, broad and
Nodule A small knoblike outgrowth, as those notched at the tip; heart shaped but attached at
found on the roots of many leguminous, con- the pointed end.
taining Rhizobium bacteria which fixes nitro- Obdeltate Inversely deltate; deltate with the
gen in the soil. broadest part at the apex.
Nom. ambig. Nomen ambiguum (Latin) ambig- Oblanceolate Inversely lanceolate, lance
uous name used in different senses which has shaped but broadest above the middle and
become a long-persistent source of error. tapering towards the base as of leaf.
Nom. cons. Nomen nonservandum (Latin) Oblate Having the shape of a spheroid with
name conserved in International Code of the equatorial diameter greater than the polar
Botanical Nomenclature. diameter; being flattened at the poles.
Nom. dub. Nomen dubium (Latin) an invalid Oblong Longer than broad with sides nearly
proposed taxonomic name because it is not parallel to each other.
accompanied by a definition or description of Obovate Inversely ovate, broadest above the
the taxon to which it applies. middle.
Nom. illeg. Nomen illegitimum (Latin) ille- Obpyramidal Resembling a four-sided pyra-
gitimate taxon deemed as superfluous at its mid attached at the apex with the square base
time of publication either because the taxon facing away from the attachment.
Scientific Glossary 369
Obpyriform Inversely pyriform, resembling a Ovule The young, immature seed in the ovary
pear which is attached at the narrower end. cf. which becomes a seed after fertilisation. adj.
pyriform. ovular.
Obspathulate Inversely spathulate; resem- Ovulode A sterile reduced ovule borne on the
bling a spoon but attached at the broadest end. placenta, commonly occurring in Myrtaceae.
cf. spathulate. Oxisols Refer to ferralsols.
Obtriangular Inversely triangular; triangular Pachymorphic Describes the short, thick rhi-
but attached at the apex. cf. triangular. zomes of clumping bamboos with short, thick
Obtrullate Inversely trullate; resembling a and solid internode (except the bud-bearing
trowel blade with the broadest axis above the internodes, which are more elongated). cf.
middle. cf. trullate. sympodial.
Obtuse With a blunt or rounded tip, the con- Palate (Botany) a raised appendage on the
verging edges separated by an angle greater lower lip of a corolla which partially or com-
than 90°. pletely closes the throat.
-oid Suffix denoting a three-dimensional shape, Palea The upper of the two membraneous
e.g. spheroid. bracts of a grass floret, usually enclosing the
Ochraceous A dull yellow colour. lodicules, stamens and ovary. pl. paleae. adj.
Ocreate Having a tube-like covering around paleal. cf. lemma.
some stems, formed of the united stipules; Paleate Having glumes.
sheathed. Palm heart Refers to soft, tender inner core
Oleaginous Oily. and growing bud of certain palm trees which
Oligotrophic Lacking in plant nutrients and are eaten as vegetables. Also called heart of
having a large amount of dissolved oxygen palm, palmito, burglar’s thigh, chonta or
throughout. swamp cabbage.
Operculum A lid or cover that becomes Palmate Describing a leaf which is divided
detached at maturity by abscission, e.g. in into several lobes or leaflets which arise from
Eucalyptus, also a cap or lid covering the bud the same point. adv. palmately.
and formed by fusion or cohesion of sepals Palmito See Palm heart.
and/or petals. adj. operculate. Palustrial Paludal, swampy, marshy.
Opposite Describing leaves or other organs Palustrine Marshy, swampy.
which are borne at the same level but on oppo- Palustrine herb Vegetation that is rooted
site sides of the stem. cf. alternate. below water but grows above the surface in
Orbicular Of circular outline, disc-like. wetland system.
Order A taxonomic rank between class and Panduriform Fiddle shaped, usually with ref-
family used in the classification of organisms, erence to two dimensions.
i.e. a group of families believed to be closely Panicle A compound, indeterminate, racemose
related. inflorescence in which the main axis bears lat-
Orifice An opening or aperture. eral racemes or spikes. adj. paniculate.
Organosols Soils not regularly inundated by Pantropical Distributed throughout the tropics.
marine waters and containing a specific thick- Papilionaceous Butterfly-like, said of the pea
ness of organic materials within the upper part flower or flowers of Papilionaceae, flowers
of the profile. which are zygomorphic with imbricate pet-
Orth. var. Orthographic variant, i.e. an incor- als, one broad upper one, two narrower lateral
rect alternate spelling of a name. ones and two narrower lower ones.
Ovary The female part of the pistil of a flower Papilla A small, superficial protuberance on
which contains the ovules (immature seeds). the surface of an organ being an outgrowth
Ovate Egg shaped, usually with reference to of one epidermal cell. pl. papillae. adj.
two dimensions. papillose.
Ovoid Egg shaped, usually with reference to Papillate Having papillae.
three dimensions. Papillose Covered with papillae.
370 Scientific Glossary
Pappus A tuft (or ring) of hairs, bristles or Perianth The two outer floral whorls of the
scales borne above the ovary and outside the angiosperm flower; commonly used when the
corolla as in Asteraceae often persisting as a calyx and the corolla are not readily distin-
tuft of hairs on a fruit. adj. pappose, pappous. guishable (as in monocotyledons).
Papyraceous Resembling parchment of paper. Pericarp (Botany). The wall of a ripened
Parenchyma Undifferentiated plant tissue ovary; fruit wall composed of the exocarp,
composed of more or less uniform cells. mesocarp and endocarp.
Parietal Describes the attachment of ovules to Persistent Remaining attached; not falling off.
the outer walls of the ovaries. cf. caduceus.
Paripinnate Pinnate with an even number Petal Free segment of the corolla. adj. petaline.
of leaflets and without a terminal leaflet. cf. cf. lobe.
imparipinnate. Petiolar Relating to the petiole.
-partite Divided almost to the base into seg- Petiolate Having petiole.
ments, the number of segments written as a Petiole Leaf stalk. adj. petiolate.
prefix. Petiolulate Supported by its own petiolule.
Patelliform Shaped like a limpet shell; cap Petiolule The stalk of a leaflet in a compound
shaped and without whorls. leaf. adj. petiolulate.
Patent Diverging from the axis almost at right pH Is a measure of the acidity or basicity of a
angles. solution. It is defined as the cologarithm of the
Peat Is an accumulation of partially decayed activity of dissolved hydrogen ions (H+).
vegetation matter. Phenology The study of periodic plant life
Pectin A group of water-soluble colloidal car- cycle events as influenced by seasonal and
bohydrates of high molecular weight found in interannual variations in climate.
certain ripe fruits. Phyllary A bract of the involucre of a compos-
Pectinate Pinnatifid with narrow segments ite plant, term for one of the scalelike bracts
resembling the teeth of a comb. beneath the flower head in Asteraceae.
Pedicel The stalk of the flower or stalk of a Phylloclade A flattened, photosynthetic branch
spikelet in Poaceae. adj. pedicellate. or stem that resembles or performs the func-
Pedicellate Having pedicel. tion of a leaf, with the true leaves represented
Peduncle A stalk supporting an inflorescence. by scales.
adj. pedunculate Phyllode A petiole that functions as a leaf. adj.
Pellucid Allowing the passage of light; trans- phyllodineous. cf. cladode.
parent or translucent. Phyllopodia Refer to the reduced, scalelike
Pellucid dotted Copiously dotted with leaves found on the outermost portion of the
immersed, pellucid, resinous glands. corm where they seem to persist longer than
Peltate With the petiole attached to the lower typical sporophylls as in the fern Isoetes.
surface of the leaf blade. Phytoremediation Describes the treatment
Pendant Hanging down. of environmental problems (bioremediation)
Pendulous Drooping, as of ovules. through the use of plants which mitigate the
Penniveined or penninerved Pinnately environmental problem without the need to
veined. excavate the contaminant material and dispose
Pentamerous In five parts. of it elsewhere.
Perennial A plant that completes its life cycle Pileus (Botany) cap of mushroom.
or lives for more than two years. cf. annual, Piliferous (Botany) bearing or producing hairs,
biennial. as of an organ with the apex having long, hair-
Perfoliate A leaf with the basal lobes united like extensions.
around—and apparently pierced by—the Pilose Covered with fine soft hairs.
stem. Pinna A primary division of the blade of a
Pergamentaceous Parchment-like. compound leaf or frond. pl. pinnae.
Scientific Glossary 371
Pinnate Bearing leaflets on each side of a Polygamous With unisexual and bisexual flow-
central axis of a compound leaf; divided into ers on the same or on different individuals of
pinnae. the same species.
Pinnatifid, pinnatilobed A pinnate leaf parted Polymorphic With different morphological
approximately halfway to midrib; when variants.
divided to almost to the midrib described as Polypetalous (Botany) having a corolla com-
deeply pinnatifid or pinnatisect. posed of distinct, separable petals.
Pinnatisect Lobed or divided almost to the Pome A fleshy fruit where the succulent tissues
midrib. are developed from the receptacle.
Pinnule A leaflet of a bipinnate compound leaf. Pore A tiny opening.
Pistil Female part of the flower comprising the Premorse Abruptly truncated, as though bitten
ovary, style and stigma. or broken off as of a leaf.
Pistillate Having one or more pistils; having Procumbent Trailing or spreading along the
pistils but no stamens. ground but not rooting at the nodes, referring
Placenta The region within the ovary to which to stems. cf. ascending, decumbent, erect.
ovules are attached. pl. placentae. Pro hyb. (Latin) as a hybrid.
Placentation The arrangement of the placentae Pro parte (Latin) in part
and ovules in the ovary. Pro parte majore (Latin) for the greater part.
Plano- A prefix meaning level or flat. Pro parte minore (Latin) for a small part.
Pleonanthic Refer to palms in which the stem Pro sp. (Latin) as a species.
does not die after flowering. Pro subsp. (Latin) as a subspecies.
Plicate Folded like a fan. Pro syn. (Latin) as a synonym.
Plumose Feather-like, with fine hairs arising Prophyll A plant structure that resembles a
laterally from a central axis; feathery. leaf.
Pneumatophore Modified root which allows Prostrate Lying flat on the ground.
gaseous exchange in mud-dwelling shrubs, Protandrous Relating to a flower in which the
e.g. mangroves. anthers release their pollen before the stigma
Pod A dry one- to many-seeded dehiscent of the same flower becomes receptive.
fruit, as applied to the fruit of Fabaceae, Proximal End of any structure closest to the
i.e. Caesalpiniaceae, Mimosaceae and point of attachment. cf. distal.
Papilionaceae. Pruinose Having a thick, waxy, powdery coat-
Podzol, podsolic soil Any of a group of acidic, ing or bloom.
zonal soils having a leached, light-coloured, Pseudocarp A false fruit, largely made up of tis-
grey and ashy appearance. Also called sue that is not derived from the ovary but from
spodosol. floral parts such as the receptacle and calyx.
Pollen cone Male cone or microstrobilus or Pseudostem The false, herbaceous stem of a
pollen cone is structurally similar across all banana plant composed of overlapping leaf bases.
conifers; extending out from a central axis are Pteridophyte A vascular plant which repro-
microsporophylls (modified leaves). Under duces by spores; the ferns and fern allies.
each microsporophyll is one or several micro- Puberulent Covered with minute hairs or very
sporangia (pollen sacs). fine down; finely pubescent.
Pollinia The paired, waxy pollen masses of Puberulous Covered with a minute down.
flowers of orchids and milkweeds. Pubescent Covered with short, soft hairs.
Polyandrous (Botany) having an indefinite Pulvinate Having a swell, pulvinus at the base
number of stamens. as a leaf stalk.
Polyembryonic seed Seeds contain many Pulvinus Swelling at the base of leaf stalk.
embryos, most of which are asexual (nucel- Pulviniform Swelling or bulging.
lar) in origin and genetically identical to the Punctate Marked with translucent dots or
maternal parent. glands.
372 Scientific Glossary
Punctiform Marked by or composed of points Retuse With a very blunt and slightly notched
or dots. apex. cf. emarginated.
Punctulate Marked with minute dots; a dimin- Revolute With the margins inrolled on the
utive of punctate. lower (abaxial) surface.
Purpurascent Purple or becoming purple. Rhizine A root-like filament or hair growing
Pusticulate Characterised by small pustules. from the stems of mosses or on lichens.
Pyrene The stone or pit of a drupe, consisting Rhizoid Root-like filaments in a moss, fern, fun-
of the hardened endocarp and seed. gus, etc., that attach the plant to the substratum.
Pyriform Pear shaped, a three-dimensional Rhizome A prostrate or underground stem con-
shape; attached at the broader end. cf. sisting of a series of nodes and internodes with
obpyriform. adventitious roots and which generally grows
Pyxidium Seed capsule having a circular lid horizontally.
(operculum) which falls off to release the seed. Rhizophore A stilt-like outgrowth of the stem
Raceme An indeterminate inflorescence with a which branches into roots on contact with the
simple, elongated axis and pedicellate flowers, substrate.
youngest at the top. adj. racemose. Rhombic Shaped like a rhombus.
Rachilla The main axis of a grass spikelet. Rhomboid Shaped like a rhombus.
Rachis The main axis of the spike or other Rib A distinct vein or linear marking, often
inflorescence of grasses or a compound leaf. raised as a linear ridge.
Radiate Arranged around a common centre; as Riparian Along the river margins, interface
of an inflorescence of Asteraceae with mar- between land and a stream.
ginal, female or neuter, ligulate ray florets and Rosette A tuft of leaves or other organs
central, perfect or functionally male, tubular, arranged spirally like petals in a rose, rang-
disc florets. cf. disciform, discoid. ing in form from a hemispherical tuft to a flat
Radical Arising from the root or its crown, or the whorl. adj. rosetted, rosulate.
part of a plant embryo that develops into a root. Rostrate Beaked; the apex tapered into a slen-
Ray The marginal portion of the inflorescence der, usually obtuse point.
of Asteraceae and Apiaceae when distinct Rostrum A beak-like extension.
from the disc. Also, the spreading branches of Rosulate Having a rosette.
a compound umbel. Rotate Wheel shaped; refers to a corolla with
Receptacle The region at the end of a pedicel a very short tube and a broad upper part
or on an axis which bears one or more flowers. which is flared at right angles to the tube. cf.
adj. receptacular. salverform.
Recurved Curved downwards or backwards. Rotundate Rounded; especially at the end or
Reflexed Bent or turned downward. ends.
Regosol Soil that is young and undeveloped, Rugae Refers to a series of ridges produced by
characterised by medium to fine-textured folding of the wall of an organ.
unconsolidated parent material that maybe Rugose Deeply wrinkled.
alluvial in origin and lacks a significant hori- Rugulose Finely wrinkled.
zon layer formation. Ruminate (Animal) chew repeatedly over an
Reniform Kidney shaped in outline. extended period.
Repand With slightly undulate margin. Ruminate endosperm Uneven endosperm sur-
Replicate Folded back, as in some corolla face that is often highly enlarged by ingrowths
lobes. or infoldings of the surrounding tissue. cf.
Resinous Producing sticky resin. homogeneous endosperm.
Resupinate Twisted through 180°. Rz value Is a numerical reference to the mesh/
Reticulate Having the appearance of a network. emulsion equalisation on the screen.
Retrorse Bent or directed downwards or back- Saccate Pouched.
wards. cf. antrorse. Sagittate Shaped like an arrow head.
Scientific Glossary 373
Saline soils Soils that contain excessive levels Secund With the flowers all turned in the same
of salts that reduce plant growth and vigour by direction.
altering water uptake and causing ion-specific Sedge A plant of the family Apiaceae,
toxicities or imbalances. Cyperaceae.
Salinity Is characterised by high electrical con- Segmented Constricted into divisions.
ductivities and low sodium ion concentrations Seminal root Or seed root originates from the
compared to calcium and magnesium scutellar node located within the seed embryo
Salverform Applies to a gamopetalous and is composed of the radicle and lateral
corolla having a slender tube and an abruptly seminal roots.
expanded limb. Senescence Refers to the biological changes
Samara An indehiscent, winged, dry fruit. which take place in plants as they age.
Sand A naturally occurring granular material Sepal Free segment of the calyx. adj. sepaline.
composed of finely divided rock and mineral Septum A partition or cross wall. pl. septa. adj.
particles range in diameter from 0.0625μm to septate.
2 mm. adj. sandy Seriate Arranged in rows.
Saponins Are plant glycosides with a distinc- Sericeous Silky; covered with close-pressed,
tive foaming characteristic. They are found in fine, straight silky hairs.
many plants, but get their name from the soap- Serrate Tooth like a saw; with regular, asym-
wort plant (Saponaria). metric teeth pointing forward.
Saprophytic Living on and deriving nourish- Serrated Toothed margin.
ment from dead organic matter. Serratures Serrated margin.
Sapwood Outer woody layer of the tree just Serrulate With minute teeth on the margin.
adjacent to and below the bark. Sessile Without a stalk.
Sarcotesta Outermost fleshy covering of Seta A bristle or stiff hair. pl. setae. adj. setose,
Cycad seeds below which is the sclerotesta. setaceous.
Scabrid Scurfy, covered with surface abra- Setaceous Bristlelike.
sions, irregular projections or delicate scales. Setate With bristles.
Scabrous Rough to the touch because of scat- Setiform Bristle shaped.
tered rough hairs. Setulose With minute bristles.
Scale Dry bract or leaf. Sheathing Clasping or enveloping the stem.
Scandent Refer to plants, climbing. Shrub A woody plant usually less than 5 m
Scape Erect flowering stem, usually leafless, rising high and with many branches without a dis-
from the crown or roots of a plant. adj. scapose. tinct main stem except at ground level.
Scapigerous With a scape. Silicula A broad, dry, usually dehiscent fruit
Scarious Dry, thin and membranous. derived from two or more carpels which
Schizocarp A dry fruit which splits into lon- usually dehisce along two sutures. cf.
gitudinally multiple parts called mericarps or siliqua.
cocci. adj. schizocarpous. Siliqua A silicula which is at least twice as
Sclerotesta The innermost fleshy coating of long as broad.
cycad seeds, usually located directly below Silt Is soil- or rock-derived granular material
the sarcotesta. of a grain size between sand and clay, grain
Scorpoid Refers to a cymose inflorescence in particles ranging from 0.004 to 0.06 mm in
which the main axis appears to coil. diameter. adj. silty.
Scutellum (Botany) any of various parts Simple Refer to a leaf or other structure that is
shaped like a shield. not divided into parts. cf. compound.
Scutiform Shaped like a shield. Sinuate With deep wavy margin.
Secondary venation Arrangement of the lat- Sinuous Wavy.
eral veins arising from the midrib in the leaf Sinus An opening or groove, as occurs between
lamina. the bases of two petals.
374 Scientific Glossary
Sodicity Is characterised by low electrical con- Sporangium A spore bearing structure found
ductivities and high sodium ion concentra- in ferns, fern allies and gymnosperms. pl. spo-
tions compared to calcium and magnesium. rangia. adj. sporangial.
Sodic soils Contains high levels of sodium salts Sporocarp A stalked specialised fruiting struc-
that affects soil structure, inhibits water move- ture formed from modified sporophylls, con-
ment and causes poor germination and crop taining sporangia or spores as found in ferns
establishment and plant toxicity. and fern allies.
Soil pH Is a measure of the acidity or basicity Sporophore A spore-bearing structure, espe-
of the soil. See pH. cially in fungi.
Solitary Usually refer to flowers which are Sporophyll A leaf or bract which bears or sub-
borne singly and not grouped into an inflores- tends sporangia in the fern allies, ferns and
cence or clustered. gymnosperms.
Sorocarp Fruiting body formed by some cellu- Sporophyte The spore-producing phase in the
lar slime moulds and has both stalk and spore life cycle of a plant that exhibits alternation of
mass. generations.
Sorophore Stalk bearing the sorocarp. Spreading Bending or spreading outwards and
Sorosis Fleshy multiple fruit formed from horizontally.
flowers that are crowded together on a fleshy Spur A tubular or saclike extension of the
stem, e.g. pineapple and mulberry. corolla or calyx of a flower.
Sorus A discrete aggregate of sporangia in Squama Structure shaped like a fish scale. pl.
ferns. pl. sori squamae.
Spadix Fleshy spikelike inflorescence with an Squamous Covered in scales.
unbranched, usually thickened axis and small Squarrose Having rough or spreading scale-
embedded flowers often surrounded by a like processes.
spathe. pl. spadices. Stamen The male part of a flower, consisting
Spathe A large bract ensheathing an inflores- typically of a stalk (filament) and a pollen-
cence or its peduncle. adj. spathaceous. bearing portion (anther). adj. staminal, stami-
Spatheate Like or with a spathe. nate .
Spathulate Spatula or spoon shaped; broad at Staminate Unisexual flower bearing stamens
the tip and narrowed towards the base. but no functional pistils.
Spicate Borne in or forming a spike. Staminode A sterile or abortive stamen,
Spiculate Spikelet bearing. often reduced in size and lacked anther. adj.
Spike An unbranched, indeterminate inflores- staminodial.
cence with sessile flowers or spiklets. adj. spi- Standard Refers to the adaxial petal in the
cate, spiciform. flower of Papilionaceae. cf. keel, wing.
Spikelet A small or secondary spike character- Starch A polysaccharide carbohydrate consist-
istics of the grasses and sedges and, generally, ing of a large number of glucose units joined
composed of two glumes and one or more flo- together by glycosidic bonds α-1-4 linkages.
rets. Also applied to the small spikelike inflo- Stellate Star shaped, applies to hairs.
rescence or inflorescence units commonly Stem The main axis of a plant, developed from
found in Apiaceae. the plumule of the embryo and typically bear-
Spine A stiff, sharp, pointed structure, formed ing leaves.
by modification of a plant organ. adj. spinose. Sterile Lacking any functional sexual parts
Spinescent Ending in a spine; modified to form which are capable of fertilisation and seed
a spine production.
Spinulate Covered with small spines. Stigma The sticky receptive tip of an ovary with
Spinulose With small spines over the surface. or without a style which is receptive to pollen.
Spodosol See Podzol. Stilt root A supporting root arising from the
Sporidia Asexual spores of smut fungi. stem some distance above the ground as in
Scientific Glossary 375
Tepal A segment of the perianth in a flower in Triveined Main laterals arising at the base of
which all the perianth segments are similar lamina.
in appearance and are not differentiated into Triquetrous Three edged; acutely three angled.
calyx and corolla; a sepal or petal. Trullate With the widest axis below the middle
Tetrasporangium A sporangium containing and with straight margins; ovate but margins
four haploid spores as found in some algae. straight and angled below middle, trowel,
Terete Having a circular shape when cross- angular, ovate shaped.
sectioned or a cylindrical shape that tapers at Truncate With an abruptly transverse end as if
each end. cut off.
Terminal At the apex or distal end. Tuber A stem, usually underground, enlarged
Ternate In threes as of leaf with three leaflets. as a storage organ and with minute scalelike
Testa A seed coat, outer integument of a seed. leaves and buds. adj. tuberous.
Thallus Plant body of algae, fungi and other Tubercle A wartlike protuberance. adj.
lower organisms. tuberculate.
Thyrse A dense, panicle-like inflorescence, as Tuberculate Bearing tubercles; covered with
of the lilac, in which the lateral branches ter- warty lumps.
minate in cymes. Tuberisation Formation of tubers in the soil.
Tomentose Refers to plant hairs that are bent Tuft A densely packed cluster arising from an
and matted forming a woolly coating. axis. adj. tufted.
Tomentellose Mildly tomentose. Turbinate Having the shape of a top; cone shaped,
Torus Receptacle of a flower. with the apex downwards, inversely conic.
Transpiration Evaporation of water from the Turgid Distended by water or other liquid.
plant through leaf and stem pores. Turion The tender young, scaly shoot such as
Tree That has many secondary branches sup- asparagus, developed from an underground
ported clear of the ground on a single main bud without branches or leaves.
stem or trunk. Turnery Articles made by the process of
Triangular Shaped like a triangle, three angled turning.
and three sided. Twining Winding spirally.
Tribe A category intermediate in rank between Ultisols Mineral soils with no calcareous mate-
subfamily and genus. rial, having less than 10% weatherable miner-
Trichome A hair-like outgrowth of the als in the extreme top layer of soil and with
epidermis. less the 35% base saturation throughout the
Trichotomous Divided almost equally into soil.
three parts or elements. Umbel An inflorescence of pedicellate flowers
Tridentate Three toothed or three pronged. of almost equal length arising from one point
Trifid Divided or cleft into three parts or lobes. on top of the peduncle. adj. umbellate.
Trifoliate Having three leaves. Umbellet A secondary umbel of a compound
Trifoliolate A leaf having three leaflets. umbel. cf. umbellule.
Trifurcate Having three forks or branches. Umbellule A secondary umbel of a compound
Trigonous Obtusely three angled; triangular in umbel.cf. umbellet.
cross section with plane faces. Uncinate Bent at the end like a hook; unciform.
Tripartite Consisting of three parts. Undershrub Subshrub; a small, usually
Tripinnate Relating to leaves, pinnately sparsely branched woody shrub less than 1 m
divided three times with pinnate pinnules. high. cf. shrub.
Tripliveined Main laterals arising above base Undulate With an edge/margin or edges wavy
of lamina. in a vertical plane; may vary from weakly to
Triploid Describing a nucleus or cell that has strongly undulate or crisped. cf. crisped.
three times (3n) the haploid number (n) of Unifoliolate A compound leaf which has been
chromosomes. reduced to a single, usually terminal leaflet.
Scientific Glossary 377
Uniform With one form, e.g. having stamens Vertosols Soils that both contain more than
of a similar length or having one kind of leaf. 35% clay and possess deep cracks wider than
cf. dimorphic. 5mm during most years.
Uniseriate Arranged in one row or at one level. Vesicle A small bladdery sac or cavity filled
Unisexual With one sex only, either bear- with air or fluid. adj. vesicular.
ing the anthers with pollen, or an ovary with Vestigial The remaining trace or remnant of
ovules, referring to a flower, inflorescence or an organ which seemingly lost all or most
individual plant. cf. bisexual. of its original function in a species through
Urceolate Shaped like a jug, urn or pitcher. evolution.
Utricle A small bladdery pericarp. Vestiture Covering; the type of hairiness, scali-
Vaginate Forming or enclosed in a sheath. ness or other covering commonly found on the
Valvate Meeting without overlapping, as of external parts of plants. cf. indumentums.
sepals or petals in bud. cf. imbricate. Vibratile Capable of to and fro motion.
Valve One of the sections or portions into Villose Covered with long, fine, soft hairs, finer
which a capsule separates when ripe. than in pilose.
Variant Any definable individual or group Villous Covered with soft, shaggy unmatted
of individuals which may or may not be hairs.
regarded as representing a formal taxon after Vine A climbing or trailing plant.
examination. Violaxanthin Is a natural xanthophyll pigment
Variegate, variegated Diverse in colour or with an orange colour found in a variety of
marked with irregular patches of different plants like pansies.
colours, blotched. Viscid Sticky, being of a consistency that
Variety A taxonomic rank below that of resists flow.
subspecies. Viviparous Describes seeds or fruit which
Vein (Botany) a strand of vascular bundle sprout before they fall from the parent
tissue. plant.
Veinlets Small veins. Whorl A ringlike arrangement of leaves,
Velum A flap of tissue covering the sporan- sepals, stamens or other organs around an
gium in the fern, Isoetes. axis.
Velutinous Having the surface covered with a Winged Having a flat, often membranous
fine and dense silky pubescence of short fine expansion or flange, e.g. on a seed, stem or
hairs; velvety. cf. sericeous one of the two lateral petals of a papiliona-
Venation Distribution or arrangement of veins ceous flower or one of the petal-like sepals of
in a leaf. Polygalaceae. cf. keel, standard.
Veneer Thin sheet of wood. Xanthophylls Are yellow, carotenoid pigments
Ventral (Botany) facing the central axis, found in plants. They are oxidised derivatives
opposed to dorsal. of carotenes.
Vernation The arrangement of young leaves or Xeromorphic Plant with special modified
fronds in a bud or at a stem apex. cf. circinnate. structure to help the plant to adapt to dry
Verrucose Warty. conditions.
Verticil A circular arrangement, as of flowers, Xerophyte A plant which naturally grows in
leaves or hairs, growing about a central point; dry regions and is often structurally modified
a whorl. to withstand dry conditions.
Verticillaster False whorl composed of a pair Zygomorphic Having only one plane of
of opposite cymes as in Lamiaceae. symmetry, usually the vertical plane,
Verticillate Whorled, arranged in one or more referring to a flower, calyx or corolla. cf.
whorls. actinomorphic.
Vertisol A soil with a high content of expansive Zygote The fist cell formed by the union of
montmorillonite clay that forms deep cracks two gametes in sexual reproduction. adj.
in drier seasons or years. zygotic.
Common Name Index
A T24, 154
A-549, 257 TCCSUP, 154
Abrupt Greenhood, 17 UMUC3, 154
Abu Suwaif, 21 Blady Grass, 22
A549 cancer cell lines, 258 Blotched Hyacinth Orchid, 11
African Potato, 4 Blue Caladenia, 9
Ahipa, 2 Blue Fairy Orchid, 9
Akar Wangi (Malay), 22 Blue Lotus Of Egypt, 7
Alang-Alang, 22 Blue Lotus Of India, 7
Ali’s umbrella, 249 Blue Water Lily, 7
Ali’s walking stick, 249 Blunt Greenhood, 18
Alliums, 2 Bodmon Sok, 4
Alpen Greenhood, 18 Boga Bhet, Seluk (Assamese), 8
Amazon Water Lily, 123 Bonmala (Assamese), 3
Amazon Water Platter, 123 Bonnet Orchid, 10
American Lotus, 6 Boorman’s Greenhood, 18
American White, 113 Breadroot, 2
Ant Orchid, 9 Breast cancer cell lines, 157
Añu, 2 Bristly Helmut Orchid, 10
A2780 ovarian tumour, 154 Broad-Lip Bird Orchid, 10
Arrow Head, 1, 48 Broad-Lipped Leek Orchid, 17
Arrowroot, 2, 5, 48, 49, 51 Brown Bird Orchid, 10
Artichoke, 2 Bullgrass, 227
Artichoke Betony, 40 Burdock, 2
Assamese, 4 Burnt-Tip Orchid, 17
Awa, 21, 146, 183 Bush potato, 2
Butterfly Orchid, 9
B
Barnyardgrass, 183 C
Baur’s Disa, Ntetemera, 12 Calcutta Cane, 21
Bearded Iris, 3, 26 Cancer cell lines, 256, 258
Beaver Root, 113 Cancerous cells (CaOv-3), 258
Bee Orchid, 15 Cao Shi Can (Chinese), 5
Beet, 2 CaOV-3 cancer cell lines, 256
Ben Lomond Leek Orchid, 17 Carrot, 2
Bermuda, 48 CaSki cervical cancer cells, 208
BGC-823, 258 Cassava, 2
Birds’s Mouth Orchid, 17 Cassia, 217
Black salsify, 2 Cedarwood, 217
Black Vetiver grass, 22 Celeriac, 2
Bladder cancer cell lines, 154 Cell lines, 257
5637, 154 CEM-SS human cancer cell line, 231
HT1197, 154 Cervical cancer HeLa, 34
HT1376, 154 Chago, 2, 6
N LAPC-4, 155
Naked Man Orchid, 15, 16 LNCaP, 155
Naked Sun Orchid, 20 PC-3, 155
Nang Guo Long Chuan Hua (Chinese), 23 22Rv1, 155
Napier grass, 217 Protozoa, 217
Ndulamo, Lisesa, 13 Provence Orchid, 16
Nepal Satyrium, 19 Pumpkin, 237
New Zealand Yam, 2, 138 Purple Donkey Orchid, 12
Nginti Ali (Mishing), 5 Purple Flag, 26
Nodding Greenhood, 18 Purple Hyacinth Orchid, 11
Northern Bugleweed, 4 Pyramidal Orchid, 9, 16
Novel, 178
Numbunumbu, 19
Q
Queen Elizabeth Root, 26
O Quiba Yam, 138
Obedience plant, St. Vincent, 48 Quick Grass, 21, 22
Oca, 2, 21, 138, 140, 141, 144 Quitch Grass, 22
Oka, 138
Onions, 2
Orach, Deppei Wood Sorrel, 20 R
Orange, 182 Radish, 2, 184
Orchid, 23 Rainforest Greenhood, 18
Orchid, Fringed spider Orchid, 9 RD, 256
Orchis De Provence, 16 Red beans, 60
Orkida kbira, 9 Red Lily of the Nile, 117
Ornate Ant Orchid, 10 Red Midge Orchid, 13
Orris, 3, 26 Red Tongue Orchid, 10
Orrisroot, 26, 27 Red Water Lily, 7, 117, 120
The Ottoman Dactylorhiza, 10 Rekang Nemthu, 23
Oval Coelogyne, 10 Rhodes grass, 214
Rizuka, 4
Rock Lily, King Orchid, 11
P Roman Dactylorhiza, 11
Packing Leaf, 6 Ronga Bhet, 8
Painted Nettle, 4 Root, 3
Papa Roja, 138 Rose, 213
Parsley, 2 Royal Water Lily, 123
Parsnips, 2 Rutabaga, 2
Patat (Sundanese), 5
Peacock flower, 3
Pelvic cancer (RD), 257 S
Persian Marsh-Orchid, 11 Sack-Carrying Dactylorhiza, 11
Phai Song Kham (Thai), 21 Saffron, 3
Phelang Riho, 4 Salsify, 2
Picroxylon Siamese, 249 Sand ginger, 2
Ping Peng Cao (Chinese), 7 Sasauge Greenhood, 19
Pink Fairies, 9 Sawfly Orchid, 15
Pink Fringe Orchid, 9 Sayabana, 4
Pink Hyacinth Orchid, 11 Scented Leek Orchid, 17
Pink Shamrock, 20 Scented Sun Orchid, 20
Pink Sun Orchid, 20 Screw-pine, 182
Pink Water Lily, 6 Shea butter, 233
Plain Sun Orchid, 20 Short-Lipped Leek Orchid, 17
Pond Collard, 7 Shui Lian (Chinese), 8
Potato Orchid, 13 Shunguneer Pushpum (Tamil), 7
Prairie turnip, 2 Siberian Motherwort, 4
Prostate cancer cell lines, 155 Sidala, 19
C4-2B, 155 SiHa, 208
DU145, 155 Silver Crabgrass, 227
Common Name Index 383
A Anatherum
Abelmochus spp, 2 A. muricatum, 196
Abronia latifolia, 6 A. zizanioides, 196
Acanthamoebae castellanii, 174 Andrographis paniculata, 262, 266
Aceras antropophorum, 8 Andropogon
Acetosella A. aromaticus, 196
A. crenata, 138 A. muricatum, 196
A.tuberosa, 138 A. muricatus, 196
Achromobacter xylosoxidans, 215 A. nardus, 196
Acianthus A. odoratus, 196
A. apprimus, 8 A. zizanioides, 196
A. collinus, 8 Anisakis simplex, 95
A. exsertus, 8 Anopheles
A. pusillus, 8 A. stephensi, 95, 211
Acinetobacter baumannii, 266–267 A. subpictus, 95
Aegle marmelos, 208 Apios americana, 2
Aframomum melegueta, 233 Arthrochilus huntianus, 9
Africana, 229 Aspergillus
Agrobacterium A. flavus, 34, 209
A. radiobacter, 111, 144 A. fumigatus, 94, 169
A. rhizogenes, 143 A. niger, 34, 94, 169
A. tumefaciens, 111 A. niger, 209
A. tume faciens, 143–144 Astragalus membranaceus, 243, 245
Agropyron Aureus, 94
A.geminatum, 227 Azadirachta indica, 35
A. repens, 21
Agrostis verticillata, 196
Akebia quinata, 80 B
Allionia expansa, 109 Bacillus
Allium sativum, 2 B. cereus, 43, 266
Alpinia galanga, 2 B.pumilus, 94
Alternaria B.subtilis, 34, 94, 209, 231, 266
A. alternata, 34 Bambusa
A. solani, 111 B.pallida, 21
Amorphallus spp., 1 B.tulda, 21
Anacamptis, 23 Barlia robertiana, 9
A. coriophora, 8, 15 Beta vulgaris, 2
A. morio, 8, 16 Bifidobacterium adolescentis, 95
A. morio subsp. picta, 9 Biomphalaria alexandrina, 35
A. palustris, 9, 16 Boerhavia
A. papilionacea, 9 B.coccinea, 6
A. pyramidalis, 9, 16 B.diffusa, 6
O P
Odorata var. stenopetala, 113 Pachyrhizus spp., 2
Olax subscorpioidea, 233 Paederia
Ondinea purpurea, 8 P. foetida, 23
Ophrys, 23 P. stenobotrya, 23
O. climacis, 14 Panax quinquefolius, 263
O. fusca, 15 Pandanus tectorius, 182
O. holosericea, 15 Pantoea agglomerans, 33
O. isaura Renz, 15 Paspalum
O. lycia Renz, 15 P. dissectum, 227
O. mammosa, 15 P. distichum, 184
O. phaseliana, 15 Pasteurella multocida, 34
O. scolopax, 15 Penicillium digitatum, 169
O. sphegodes, 15 Pennisetem purpureum, 217
Oplismenus abortivus, 196 Pennisetum clandestinum, 214
Orchis, 23 Peristylus constrictus, 17
O. adenocheila, 15 Phalaris zizanioides, 196
O. anatolica, 15 Pheretima posthuma, 120
O. anthropophora, 8, 15 Phlomis tuberosa, 4
O. conopea, 15 Phragmites, 215
O. coriophora, 15 P. australis, 22
O. italica, 15 Phragmites mauritianus, 216
O. latifolia, 16 Phrynium
O. longicruris, 16 P. capitatum, 6
O. maculata, 16 P. pubinerve, 6
O. mascula, 16 P. variegatum, 48
O. militaris, 16 Phytophthora cinnamomi, 143
O. morio, 16 Pichia stipitis, 213
O. palustris, 16 Pimedium grandiflorum, 263
O. provincialis, 16 Piper
O. pyramidalis, 16 P. methysticum, 21, 146, 147, 152, 159, 169, 170, 175
O. saccifera Brong, 16 P. wichmanii, 147
O. simia, 16 Plasmodium
O. spitzelii, 16 P. berghei, 232
O. tridentata, 17 P. falciparum, 35, 93, 258, 259
O. ustulata, 17 P. yoelii, 263
Orientalis Grueter, 20 P. yoelii-, 263
Orthoceras strictum, 17 Platanthera
Oryzias latipes, 36 P. bifolia, 17
Osmanica, 10 P. calvigera, 17
Oxalis Plectranthus, 4
O. arracacha, 138 P. barbatus, 4
O. chicligastensis, 138 P. edulis, 5
O. corymbosa, 20 P. esculenta, 2
O. crassicaulis, 138 P. esculentus, 4, 5
O. crenata, 138 P. madagascariensis, 5
O. debilis var. corymbosa, 20, 21 P. rotundifolius, 4, 5
O. deppei, 20 Poa spicata, 227
O. melilotoides var. argentina, 138 Polygonum minus, 259, 260
O. perennans, 20 Porrifolius, 2
O. picchensis, 139 Prasophyllum, 17
O. radicosa, 20 P. brevilabre, 17
O. stricta, 20 P. campestre, 17
O. tuberosa, 2, 21, 138, 139, 143 P. dossenum, 17
O. violacea, 21 P. elatum, 17
Oxalis sp., 20 P. flavum, 17
Oxybaphus expansa, 109 P. odoratum, 17
Scientific Name Index 391
P. patens, 17 Rosa
P. rogersii, 17 R. damascena, 213
P. solstitium, 17 R. laevigata, 245
Proteus vulgaris, 266 Rotundifolius, 4
Pseudomonas Rubus chingii, 245
P. aereofaciens, 144
P. aeruginosa, 43, 209, 231, 266
P. syringae, 111 S
Pseudotrichonympha grassii, 217 Saccharomyces, 213
Psoralea S. cerevisiae, 209
P. corylifolia, 245 Saccharomyces pastorianus, 169
P. esculenta, 2 Saccharum spontaneum, 22
Pteroglossaspis eustachya, 17 Sagittaria spp., 1
Pterostylis, 18 Salmonella, 34, 268
P. abrupta, 17 S. choleraesuis, 231
P. alata, 17 S. enteritidis, 52, 266
P. bicolor, 17 S. typhi, 209, 267
P. boormanii, 18 S. typhimurium, 94, 96, 174
P. chaetophora, 18 Salmonella sp., 94
P. coccina, 18 Satyrium, 23
P. curta, 18, 133 S. acutirostrum, 19
P. cycnocephala, 18 S. amblyosaccos, 19
P. daintreana, 18 S. atherstonei, 19
P. decurva, 18 S. breve, 19
P. fischii, 18 S. buchananii, 19
P. hamata, 18 S. carsonii, 19
P. hildae, 18 S. chlorocorys, 19
P. laxa, 18 S. crassicaule, 19
P. longicurva, 18 S. nepalense, 19
P. longifolia, 18 S. robustum, 19
P. longipetala, 18 S. sacculatum, 20
P. mutica, 18 S. volkensii, 20
P. nutans, 18 Scenedesmus quadricauda, 98
P. obtusa, 18 Schisandra chinensis, 245
P. parviflora, 18 Schumannianthus, 6
P. pedunculata, 18, 135, 136 Scorzonera hispanica, 2
P. praetermissa, 18 Scutellarioides, 4
P. reflexa, 18 Serapias orientalis, 20
P. setifera, 18 Serapias vomeracea ssp., 20
P. species, 19 Serrata, 4
P. truncate, 19 Serratia marcescens, 34, 144, 266
P. woollsii, 19 Shigella sp., 94
Pueraria, 237 Sium sisarum, 2
Pyricularia grisea, 184 Smallanthus sonchifolius, 2
Pythium irregulare, 111 Solenostemon rotundifolia, 5
Sorghum zizanioides, 196
Stachys
R S. affinis, 2, 5, 40–45
Rhaphis muricata, 196 S. chinensis, 5
Rhaphis zizanioides, 196 S. geobombycis var. alba, 40
Rhizoctonia solani, 143 S. pauciflora, 40
Rhizophora, 198 S. sieboldii, 5, 40, 41, 43, 44
Rhizopus S. sieboldii var. alba, 40
R. nigricans, 169 S. sieboldii var. glabrescens, 40
R. stolonifer, 184 S. sieboldii var. malacotricha, 40
Roeperocharis wentzeliana, 19 S. tuberifera, 40
Romulea bulbocodium, 3 Stachys adulterina, 5
392 Scientific Name Index
Staphylococcus, 94 Triticum
S. aureus, 34, 35, 43, 52, 209, 231, 266 T. geminatum, 227
Steveniella satyrioides, 20 T. repens, 22
Streptococcus faecalis, 209 Tropaeolum tuberosum, 2
Subspecies africana, 230 Trypanosoma brucei brucei, 169
Typha spp, 2
Typhimurium, 34
T
Tamara
T. alba, 54 U
T. hemisphaerica, 54 Ullucus tuberosus, 2
T. rubra, 54
Taphrina deformans, 184
Taraxacum mongolicum, 243 V
Tetracera alnifolia, 233 Vetiveria
Thalia geniculata, 6 V. muricata, 196
Thanatephorus cucumeris, 184 V. nemoralis, 215
Thelymitra V. nigritana, 22
T. carnea, 20 V. odorata, 196
T. circumsepta, 20 V. zizanioides, 22, 196, 201, 207, 208,
T. cyanea, 20 210, 212–216
T. fragrans, 20 V. zizanioides var. tonkinensis, 196
T. megacalyptra, 20 Victoria
T. nuda, 20 V. amazonica, 123, 124, 125
T. pauciflora, 20 V. regia, 123
Tigrida pavonia, 3 V. regia var. randii, 123
Tipularia discolour, 20 V. regina, 123
Torulopsis petrophilum, 172 Vigna lanceolata, 2
Toxoplasma gondii, 264, 265
Trachyspermum ammi., 95
Tragopogon, 2 X
Trichoderma Xanthosoma spp., 1
T. harzianum, 111 Xanthoxalis
T. reesei, 111 X. crassicaulis, 138
T. viride, 184 X. tuberosa, 138
Trichophyton, 94 Xenopus, 85
T. ferrugineum, 169
T. longifusus, 34
T. mentagrophytes, 169, 209 Z
T. rubrum, 94, 169 Zingiber spp., 2
T. tonsurans, 169 Zizaniopsis miliacea, 22