Autotrophic plants in brief

In plant kingdom there is a competition for habitat (biotop), nutrition (food), and reproduction. For autotrophic plants nutrition is guaranteed by formation of: Leaves

chlorophyll + solar energy organic substances (sugar, starch etc.) uptake of water and minerals anorganic substances

Roots

Evolution of heterotrophic plants

Competition takes place in particular in optimal habitats. Therefore, plants or even groups of plants have to move to less favorable and sparsely populated habitats. Resettlement will only be successful if food can be obtained elsewhere. Evolution is opportunistic and parasites are the greatest opportunists. Nutrition specialists are normally heterotroph, i.e. they need for their production of energy and their physical build up already processed organic material. Hence, they are directly or indirectly dependent on autotrohic plants for carbohydrates, nitrogen, water, etc.

Relationship between a host and a root parasite

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Definition of parasitism
A coexistance of two different organisms, which supports one of them (parasite) on the expense of the other (host).

History of research on parasitic flowering plants

Theophrast (371-286 B.C.): Orobanche on Vicia ervilia sp. Plini (24-70 A.C.): Mentioned Viscum spp. Dioscorides (lst century A.C.): Orobanche spp. on legumes Mathioli (16th Century A.C.): Drawings of O. crenata and O. ramosa on various crops Tournefort (1700): Description of Cuscuta spp. von Linne (1753): First description of Striga spp.

Bentham (1835): Description of 14 Striga spp. Beck von Mannagetta (19th century): Mentioned 130 Orobanche spp. Heinricher (1910-1930): Studies on 18 European plant parasites Yunker (1920): Review of Cuscuta spp. Tubeuf (1923): Monographs of Viscum spp. Saunders (1930): First comprehensive study on Striga Kuijt (1969): First comprehensive taxonomic study on all parasitic plants

Parasitic flowering plants Arab view and philosophy of the nature in the 10th century, from Dieterici (1861)
In the plant kingdom there are types of plants, which are in their behavior like the soul of animals, whereas their body is a body of plants: these are the parasitic plants. Unlike the other type of plants, parasitic plants have neither roots nor leaves as the others. However, they attach themselves to the trees, seeds and thorns from which they absorb moisture and nutrients like the worm does, who creeps on leaves of the trees and stems of plants, cuts up and eats to absorb his nutrients. Research on Orobanche started mainly in Europe already at the beginning of last century, while work on Striga began only at the beginning of this century.

Parasites in plant kingdom

The parasitic way of life has developed and established itself independently in many different families of plant kingdom. Parasitism is only known from dicotyledonous plants. Today we know about 20 families where parasitism occurs, with 3000-5000 species. Families of economic importance are as follows: Loranthaceae (70/900)* Dendrophtoe Tapinantus Amyema Viscaceae (7/500) Viscum Phoradendron Arceutobium Scrophulariaceae (10/500) Striga Rhinanthus Alectra Parentucellia Melampyrum Orobanchaceae (14/160) Orobanche Cistanche Phelypaea Cuscutaceae (1/180) Cuscuta Lauraceae (1/30) Cassytha

Pedicularis Bellardia * (Number of genera with parasites/number of parasitic species)

Definitions for the description of plant parasites

Parasite: A plant depending on another plant for part or all of its nutrition. Hemi-parasite or semi-parasite: A plant which is only partially parasitic, possessing its own chlorophyll (green colour) and photosynthetic ability (may be facultative or obligate - see below). Holo-parasite: A plant which is totally parasitic, lacking chlorophyll and thus unable to synthesize organic carbon. Obligate parasite: A plant which cannot establish and develop independently. Facultative parasite: A plant which can establish and grow independently but which normally behaves as a parasite to obtain some of its nutrition. Haustorium: The bridge of tissue connecting the host and parasite, usually a swollen mass consisting of both host and parasite tissue. this acts as a conduit for the flow of water and nutrient from host to parasite. You can find some more definitions in the "glossary" annexe to the technical manual Important characteristics in parasitic life of selected genera Alectra Lack of chlorophyll Fleshy stem Microspermy Embryo rudimentary Need for stimulation of germination Reduction of root system Primary and secondary haustoria + = yes; = no + + + + Striga + + + + + + + Cuscuta + + Orobanche + + + + + + +

Criterion of intensity of parasitism

Structures which are essential for normal plants to survive, often are not necessary for parasites, so that these structures get lost during evolution. An extreme example is Rafflesia arnoldii. This plant has thrown off all the unneeded organs with the exception of an efficient haustorial system, and a flower which is the biggest in plant kingdom.

Afterripening and conditioning of seed

Seeds of Orobanche, Striga and Alectra are dormant and require a period of afterripening in warm, dry storage, followed by conditioning in a warm, moist environment, before they will respond to germination stimulants. Cuscuta as a representative of shoot parasites does not require host stimulants.

Early development stages of the genera Alectra, Orobanche and Striga

Germination and attachment of Cuscuta The length and effectiveness of the afterripening period may vary with temperature, humidity, and other environmental factors, e.g. Striga seeds became receptive to conditioning after storage for 6 weeks at 31C. At lower temperatures, longer afterripening periods were required, e.g. 32 weeks at 4C. The nature of the changes occurring in the seeds during afterripening are unknown. Various structural and metabolic changes in the seed coat and embryo may occur, and changes in the respiratory substrate are involved in the afterripening. The dormancy in Cuscuta seed is mainly due to the seed coat itself. Cuscuta seed therefore may stay dormant for several years upon the testa is rotted; on the other hand, part of the seeds may germinate directly after maturity on the mother plant. Before the seeds of parasites will respond to germination stimulants, they require a period of conditioning subsequent to afterripening. Seed coat permeability may increase during conditioning and/or changes in the levels or activities of endogenous germination promoters or inhibitors may occur. The duration of conditioning is dependent on certain factors like temperature, origin, and age of the seeds. The period of time necessary for conditioning ranges from several days up to several weeks. Optimal temperature during conditioning

Species Striga hermonthica Striga asiatica Striga gesnerioides Orobanche ramosa Orobanche crenata Alectra vogelii

Temperature 23C 30C 23C 20-25C 18C 23C

Prolonged conditioning without subsequent germination induction by stimulants can cause secondary dormancy ("wet dormancy").

Germination
Seeds of the parasites in the dry state can remain dormant and viable for many years. Germination in the field starts about 10 days after the beginning of the rainy season when the host plant has already established. In the laboratory, a water-conditioning period of 6 to 10 days readies the seed for germination. Striga will germinate in 1 to 2 days, Orobanche in 5 to 7 days after exposure to the host root exudate. With Cuscuta, soaking the seeds for 30 minutes in concentrated sulfuric acid will ensure high germination. The germtube elongates in the absence of a host root until the food reserves of the seed are depleted. If the germtube is close to the host root, surface elongation stops abruptly. The parasite starts to attach and to penetrate the host root epidermis. Only the seeds which receive a chemical stimulant produced by the host roots will germinate. The prerequisite of such a signal ensures that a host is available and near enough to be reached by a germtube which grows in direction of the origin of the signal (chemotropism). Most of the seed population won't be reached by the stimulant and remain viable in the soil until the following cropping periods. The germination stimulant is exuded 3-6 mm behind the root tip of the hostplant. No stimulant is produced in older root cells. Germination stimulants of different hosts can have a synergistic effect, e.g. maize and flax alone stimulated 4% to 5% of the Striga seeds tested. When the root exudates of both hosts were mixed together, germination of Striga seeds was stimulated up to 51%. Germination stimulants can be replaced by several other substances like kinetin, giberellic acid, abscisic acid, cumarin derivates, yeast extracts, ethylene etc. This demonstrates that for stimulating germination of parasitic seeds not necessarily a host specific stimulant is required.

Development of a haustorium by a parasite Roots of different nonhost plants are able to produce germination stimulants for parasites as well. But attachment itself is normally restricted only to a very narrow host range. Even in one parasite species, races occur, which only attack specific host plants. These strains are reported from West Africa and India for Striga hermonthica and S. asiatica on sorghum and millet. These observations are of special interest for resistance breeding. The occurrence of Orobanche cumana strains with different degrees of aggressiveness is reported from Russia and Spain. In Cuscuta spp. seeds do not need any stimulation by the host. Germination occurs (permeability of seed testa provided) when temperature and moisture conditions are favorable, even immediately after harvest. Upon germination, the epicotyl (3-6 cm long) 'searches' for a host by circumnutation; it is even able to move for some distance in order to reach a host and to establish contact to it.

Host recognition
Cuscuta spp. contain enough food reserves in their seeds for 4 to 9 days while they are 'searching' for a suitable host. Host recognition is developed poorly as the Cuscuta stems coil and twine on almost everything. Having contacted a suitable host, haustoria from the parasite soon penetrate it, and a parasitic connection is rapidly established. Seeds of mistletoes are distributed by birds following a certain pattern (Viscum spp.). Or they are equipped with explosive fruits like Arceuthobium, which shoot them upto 15 m wide. Another possibility of host recognition developed during evolution in Orobanche and Striga is the depency of germination on germination stimulants which are released from host roots.

Haustorium
The haustorium is the salient feature of parasitic plants. It is the morphological and physiological bridge between host and parasite. Haustorium originally is defined as that part of the parasite, that develops within the tissue of the host.

The primary and secondary haustoria of the parasite and a host root Actually, the bottleneck in parasite development is the haustorium induction. For Striga a second signal is necessary to induce haustorium formation. Many parasitic plants are characterized by a poorly developed root system. Root function is augmented by haustoria which attach themselves to the root of the host plant, and penetrate it in order to absorb water, mineral nutrients and organic materials. Soon after the germination

of the parasite seed the radicle tip becomes transformed, penetrates, and attaches to the host root. Penetration of the haustorial cells into host tissue is carried out mechanically by pressure on the host endodermal cells and by hydrolytic enzymes. In some cases, this socalled primary haustorium remains the only connection with the host plant, and ensures a flow of nutrients into the parasite. Usually, the primary haustorium is supplemented by a large number of secondary haustoria which arise laterally from the parasite roots. This results in an extensive and intricate system of points of attachment between parasite and host. Secondary haustoria are not restricted to the roots of one host, but different host plants of the same species can be attached (=intraspecific parasitism), or even different host species may be parasitized simultaneously (= interspecific parasitism). A very well developed xylem and phloem continuity between parasite and host exists for Orobanche, while for Striga phloem seems to be lacking. Since Striga is holoparasitic in its subterranean stages and consequently entirely host dependent, the question is, how it obtains phloem-translocated substances such as carbohydrates? Studies on the primary haustorium of the closely related Alectra vogelli concluded, that host and parasite phloem is separated by a few layers of parenchymatous cells acting possibly as a phloem bridge. The primary functions of the haustorium is the procurement of nutrients. It should be considered, therefore, as an evolutionary adaptation for nutrient uptake. To maintain a continued influx of water into the parasite, an osmotic gradient is required. Potassium seems to be the preferred osmoticum in certain parasitic relationships. The strength with which water is drawn from the host becomes apparent under water stress, where the host shows symptoms of wilting, long before the parasite is visibly affected. Movement of solutes and other metabolites is mainly undirectional, i.e. into the parasite, e.g. phosphorus is retained by the parasite despite the fact that the host may even suffer severe deficiencies of the particular essential nutrient.

C) Classification based on dependence on other hosts: a. Stem parasite: Amerbel b. Root parasite: Striga on Jowar, Sugarcane, and Bambakhu on Tobacco, Brinjal or Chilli. c. Independent: Chandvel.