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Red algae

The red algae, or Rhodophyta (/rodft/ roh-DOFt-t or /rodfat/ ROH-d-FY-t; from Ancient
Greek: rhodon, rose and phyton,
plant), are one of the oldest groups of eukaryotic
algae,[2] and also one of the largest, with about 5,000
6,000 species[3] of mostly multicellular, marine algae, including many notable seaweeds. Other references indicate as many as 10,000 species;[4] more detailed counts
indicate about 4,000 in about 600 genera (3,738 marine
species in 546 genera in 10 orders, plus the unclassiable; and 164 freshwater species in 30 genera in eight
orders).[5]

from the Cambrian period. Other algae of dierent origins lled a similar role in the late Paleozoic, and in more
recent reefs.
Calcite crusts that have been interpreted as the remains of
coralline red algae, date to the terminal Proterozoic.[10]
Thallophytes resembling coralline red algae are known
from the late Proterozoic Doushantuo formation.[11]

3 Taxonomy

The red algae form a distinct group characterized by


having eukaryotic cells without agella and centrioles,
chloroplasts that lack external endoplasmic reticulum,
contain unstacked thylakoids[4] and use phycobiliproteins
as accessory pigments (giving them their red color). They
store oridean starch, a type of starch that consists of
highly branched amylopectin without amylose,[6] as food
reserves outside their plastids. Most red algae are also
multicellular, macroscopic, marine, and use sexual reproduction to reproduce. They have alternation of generations and may have three generations rather than two.[7]

Further information: Wikispecies:Rhodophyta


In the system of Adl et al. 2005, the red algae are classied in the Archaeplastida, along with the glaucophytes
and green algae plus land plants (Viridiplantae or Chloroplastida). The authors use a hierarchical arrangement
where the clade names do not signify rank; the class name
Rhodophyceae is used for the red algae. No subdivisions
are given; the authors say, Traditional subgroups are articial constructs, and no longer valid.[12]

The coralline algae, which secrete calcium carbonate and


play a major role in building coral reefs, belong here.
Red algae such as dulse (Palmaria palmata) and laver
(nori/gim) are a traditional part of European and Asian
cuisines and are used to make other products such as agar,
carrageenans and other food additives.[8]

The system reected the consensus in 2005. Many studies published since then have provided evidence that is in
agreement.[13][14][15][16] However, other studies have suggested Archaeplastida is paraphyletic.[17][18] As of January 2011, the situation appears unresolved.
Below are other published taxonomies of the red algae,
although none necessarily has to be used, as the taxonomy
of the algae is still in a state of ux (with classication
above the level of order having received little scientic
attention for most of the 20th century).[19]

Habitat

Most rhodophytes are marine, although freshwater


species are found; these generally prefer clean, running
water, but with some exceptions.[9]

If one denes the kingdom Plantae to mean the Archaeplastida, the red algae will be part of that kingdom

Fossil record

If Plantae are dened more narrowly, to be the


Viridiplantae, then the red algae might be considered their own kingdom, or part of the kingdom
Protista.

One of the oldest fossils identied as a red alga is also


the oldest fossil eukaryote that belongs to a specic modern taxon. Bangiomorpha pubescens, a multicellular fossil
from arctic Canada, strongly resembles the modern red
alga Bangia despite occurring in rocks dating to 1.2 bil- A major research initiative to reconstruct the Red Algal
lion years ago.[1]
Tree of Life (RedToL) using phylogenetic and genomic
Red algae are important builders of limestone reefs. The approaches is funded by the National Science Foundation
earliest such coralline algae, the solenopores, are known as part of the Assembling the Tree of Life Program.
1

3.1

PIT CONNECTIONS AND PIT PLUGS

6 Relationship to Chromalveolata
chloroplasts

Classication comparison

Some sources (such as Lee) place all red algae into the
class Rhodophyceae. (Lees organization is not a comprehensive classication, but a selection of orders consid- Chromalveolatas seem to have evolved from Bikonts that
have acquired red algae as endosymbionts. According to
ered common or important.[21] )
this theory, over time these Bikonts and their endosymSee also: Eukaryote Phylogeny
biont red algae have evolved to become Chromalveolata and their chloroplasts. This part of endosymbiotic
theory is supported by various structural and genetic
similarities.[28]

Species of red algae

Around 6,500 to 10,000 species are known,[4][8] nearly 7 Chemistry


all of which are marine, with about 200 that live only in
fresh water. However, estimates of the number of real The 13 C values of red algae reect their lifestyles.
species vary by 100%.[4]
The largest dierence results from their photosynthetic
metabolic pathway: algae that use HCO3 as a carbon
Some examples of species and genera of red algae are:
source have far more negative 13 C values than those
that only use CO2 .[29] An additional dierence of about
Cyanidioschyzon merolae, a primitive red alga
1.71 separates groups intertidal from those below the
lowest tide line, which are never exposed to atmospheric
Atractophora hypnoides
carbon. The latter group uses the more 13 C-negative CO2
dissolved in sea water, whereas those with access to atmo Gelidiella calcicola
spheric carbon reect the more positive signature of this
reserve.
Lemanea, a freshwater genus
Red algae are red due to phycoerythrin. They contain
the sulfated polysaccharide carrageenan in the amorphous
sections of their cell walls, although red algae from the
genus Porphyra contain porphyran. They also produce a
specic type of tannin called phlorotannins, but in lower
amount than brown algae do.

Palmaria palmata, dulse


Schmitzia hiscockiana
Chondrus crispus, Irish moss
Mastocarpus stellatus
Vanvoorstia bennettiana, became extinct in the early
20th century

8 Morphology

Red algae have double cell walls.[30] The outer layers contain the polysaccharides agarose and agaropectin that can
Audouinella, with freshwater as well as marine be extracted from the cell walls by boiling as agar.[30] The
internal walls are mostly cellulose.[30]
species
Acrochaetium eorescens

9 Pit connections and pit plugs

Genomes of red algae

Complete genome sequences are only available for 5 Main article: Pit connection
species of red algae, including 4 published in 2013.
Cyanidioschyzon merolae, Cyanidiophyceae[22][23]
Galdieria sulphuraria, Cyanidiophyceae

9.1 Pit connections

[24]

Pyropia yezoensis, Bangiophyceae

[25]

Chondrus crispus, Florideophyceae[26]


Porphyridium purpureum, Porphyridiophyceae[27]

Pit connections and pit plugs are unique and distinctive


features of red algae that form during the process of
cytokinesis following mitosis.[31][32] In red algae, cytokinesis is incomplete. Typically, a small pore is left in the
middle of the newly formed partition. The pit connection
is formed where the daughter cells remain in contact.

10.2

Life cycle

Shortly after the pit connection is formed, cytoplasmic The polyamine spermine is produced, which triggers carcontinuity is blocked by the generation of a pit plug, pospore production.[2]
which is deposited in the wall gap that connects the cells. Spermatangia may have long, delicate appendages, which
Connections between cells having a common parent cell increase their chances of hooking up.[2]
are called primary pit connections. Because apical growth
is the norm in red algae, most cells have two primary pit
10.2 Life cycle
connections, one to each adjacent cell.
Connections that exist between cells not sharing a common parent cell are labeled secondary pit connections.
These connections are formed when an unequal cell division produced a nucleated daughter cell that then fuses
to an adjacent cell. Patterns of secondary pit connections
can be seen in the order Ceramiales.[32]

9.2

Pit plugs

After a pit connection is formed, tubular membranes appear. A granular protein, called the plug core, then forms
around the membranes. The tubular membranes eventually disappear. While some orders of red algae simply
have a plug core, others have an associated membrane at
each side of the protein mass, called cap membranes. The
pit plug continues to exist between the cells until one of
the cells dies. When this happens, the living cell produces
a layer of wall material that seals o the plug.

9.3

Function

They display alternation of generations; in addition to


gametophyte generation, many have two sporophyte generations, the carposporophyte-producing carpospores,
which germinate into a tetrasporophyte this produces spore tetrads, which dissociate and germinate into
gametophytes.[2] The gametophyte is typically (but not
always) identical to the tetrasporophyte.[34]
Carpospores may also germinate directly into thalloid
gametophytes, or the carposporophytes may produce
a tetraspore without going through a (free-living)
tetrasporophyte phase.[34] Tetrasporangia may be arranged in a row (zonate), in a cross (cruciate), or in a
tetrad.[2]
The carposporophyte may be enclosed within the gametophyte, which may cover it with branches to form a
cystocarp.[34]
These case studies may be helpful to understand some of
the life histories algae may display:
In a simple case, such as Rhodochorton investiens:

In the Carposporophyte: a spermatium merges with a trichogyne (a long hair on the female sexual organ), which
The pit connections have been suggested to function as
then divides to form carposporangia which produce carstructural reinforcement, or as avenues for cell-to-cell
pospores.
communication and transport in red algae, however lit[33]
Carpospores germinate into gametophytes, which protle data supports this hypothesis.
duce sporophytes. Both of these are very similar; they
produce monospores from monosporangia just below a
cross wall in a lament[2] and their spores are liberated
10 Reproduction
through apex of sporangial cell.[2]
The reproductive cycle of red algae may be triggered by The spores of a sporophyte produce either tetrasporophytes. Monospores produced by this phase germinate
factors such as day length.[2]
immediately, with no resting phase, to form an identical copy of parent. Tetrasporophytes may also produce a carpospore, which germinates to form another
10.1 Fertilization
tetrasporophyte.[2]
Red algae lack motile sperm. Hence, they rely on water currents to transport their gametes to the female organs although their sperm are capable of gliding to a
carpogonium's trichogyne.[2]
The trichogyne will continue to grow until it encounters
a spermatium; once it has been fertilized, the cell wall at
its base progressively thickens, separating it from the rest
of the carpogonium at its base.[2]

The gametophyte may replicate using monospores, but


produces sperm in spermatangia, and eggs"(?) in
carpogonium.[2]
A rather dierent example is Porphyra gardneri:

In its diploid phase, a carpospore can germinate to


form a lamentous conchocelis stage, which can also
self-replicate using monospores. The conchocelis stage
eventually produces conchosporangia. The resulting
Upon their collision, the walls of the spermatium and car- conchospore germinates to form a tiny prothallus with
pogonium dissolve. The male nucleus divides and moves rhizoids, which develops to a cm-scale leafy thallus. This
into the carpogonium; one half of the nucleus merges with too can reproduce via monospores, which are produced
the carpogoniums nucleus.[2]
inside the thallus itself.[2] They can also reproduce via

13

REFERENCES

spermatia, produced internally, which are released to


meet a prospective carpogonium in its conceptacle.[2]

[4] W. J. Woelkerling (1990). An introduction. In K. M.


Cole; R. G. Sheath. Biology of the Red Algae. Cambridge
University Press, Cambridge. pp. 16. ISBN 0-52134301-1.

11

[5] Dixon, Peter S. (1977). Biology of the Rhodophyta


(Reprint. ed.). Koenigstein: Koeltz. ISBN 0-05-002485X.

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[6] Viola, R.; Nyvall, P.; Pedersn, M. (2001). The unique


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Some red algae are iridescent when not covered with water

[9] Eloranta, P.; Kwandrans, J. (2004). Indicator value of


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[10] Grant, S. W. F.; Knoll, A. H.; Germs, G. J. B. (1991).

Several species are important food crops, in particular


Probable Calcied Metaphytes in the Latest Proterozoic
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Nama Group, Namibia: Origin, Diagenesis, and Implica(Japan), gim (Korea), or laver (Britain). Dulse (Palmaria
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in Japan goes back more than three centuries.
In East and Southeast Asia, agar is most commonly produced from Gelidium amansii.

12

See also

Brown algae
Green algae
History of phycology

13

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14 External links
AlgaeBase: Rhodophyta
Seaweed Site: Rhodophyta
Tree of Life: Rhodophyta
Monterey Bay Flora

15

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