Red algae

The red algae, or Rhodophyta (/roʊˈdɒfɪtə/ roh-DOF-it-ə, /ˌroʊdəˈfaɪtə/ ROH-də-FY-tə; from Ancient Greek ῥόδον (rhodon), meaning 'rose', and
Red algae
φυτόν (phyton), meaning 'plant'), are one of the oldest groups of eukaryotic algae.[2] The Rhodophyta also comprises one of the largest phyla of
algae, containing over 7,000 currently recognized species with taxonomic revisions ongoing.[3] The majority of species (6,793) are found in the Temporal range:
[1]
Florideophyceae (class), and mostly consist of multicellular, marine algae, including many notable seaweeds.[3][4] Approximately 5% of the red Mesoproterozoic–present
[5] There are no terrestrial species, which is assumed to
algae occur in freshwater environments with greater concentrations found in the warmer area. Had'n Archean Proterozoic Pha.
be traced back to an evolutionary bottleneck where the last common ancestor lost about 25% of its core genes and much of its evolutionary
plasticity.[6]

The red algae form a distinct group characterized by having eukaryotic cells without flagella and centrioles, chloroplasts that lack external
endoplasmic reticulum and contain unstacked (stoma) thylakoids, and use phycobiliproteins as accessory pigments, which give them their red
color.[7] Red algae store sugars as floridean starch, which is a type of starch that consists of highly branched amylopectin without amylose,[8] as
food reserves outside their plastids. Most red algae are also multicellular, macroscopic, marine, and reproduce sexually. The red algal life history is
typically an alternation of generationsthat may have three generations rather than two.[9]

Chloroplasts evolved following an endosymbiotic event between an ancestral, photosynthetic cyanobacterium and an early eukarytoic
phagotroph.[10] This event (termed primary endosymbiosis) resulted in the origin of the red and green algae, and the glaucophytes, which make up
the oldest evolutionary lineages of photosynthetic eukaryotes.[11] A secondary endosymbiosis event involving an ancestral red alga and a
heterotrophic eukaryote resulted in the evolution and diversification of several other photosynthetic lineages such as Cryptophyta, Haptophyta,
Stramenopiles (or Heterokontophyta), Alveolata, Centrohelids, Katablepharids, and Telonemi.[11][12]
A-D : Chondrus crispus Stackhouse,
The coralline algae, which secrete calcium carbonate and play a major role in building coral reefs, belong here. Red algae such as dulse (Palmaria
E-F : Mastocarpus stellatus J.Ag.
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.[13] Scientific classification
Domain: Eukaryota
Unicellular members of the Cyanidiophyceae are thermoacidophiles and are found in sulphuric hot springs and other acidic environments.[14] The
remaining taxa are found in marine and freshwater environments. Most rhodophytes are marine with a worldwide distribution, and are often found at (unranked): Diaphoretickes
greater depths compared to other seaweeds because of dominance in certain pigments (i.e., phycoerythrin) within their chloroplasts.[15] Some (unranked): Archaeplastida
marine species are found on sandy shores, while most others can be found attached to rocky substrata.[15] Freshwater species account for 5% of red
algal diversity, but they also have a worldwide distribution in various habitats;[5] they generally prefer clean, high-flow streams with clear waters
Division: Rhodophyta
and rocky bottoms, but with some exceptions.[16] A few freshwater species are found in black waters with sandy bottoms [17] and even fewer are Wettstein, 1922
found in more lentic waters.[18] Both marine and freshwater taxa are represented by free-living macroalgal forms and smaller endo/epiphytic/zoic Classification is currently
forms, meaning they live in or on other algae, plants, and animals [7] In addition, some marine species have adopted a parasitic lifestyle and may be disputed. See Taxonomy.
found on closely or more distantly related red algal hosts[19][20]

Contents
Taxonomy
Classification comparison
Species of red algae
Morphology
Pit connections and pit plugs
Pit connections
Pit plugs
Function

Reproduction
Fertilization
Life cycle
Chemistry
Genomes of red algae
Fossil record
Relationship to other algae
Human consumption
Gallery
See also
References
External links

Taxonomy
In the system of Adl et al. 2005, the red algae are classified 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
artificial constructs, and no longer valid."[21]

[22][23][24][25] However, other studies have suggested

Many studies published since Adlet al. 2005 have provided evidence that is in agreement for monophyly in the Archaeplastida (including red algae).
Archaeplastida is paraphyletic.[26][27] As of January 2011, the situation appears unresolved.

Below are other published taxonomies of the red algae using molecular and traditional alpha taxonomic data; however
, the taxonomy of the red algae is still in a state of flux (with classification above the
[28]
level of order having received little scientific attention for most of the 20th century).

If one defines the kingdom Plantae to mean the Archaeplastida, the red algae will be part of that kingdom
 If Plantae are defined more narrowly, to be the Viridiplantae, then the red algae might be considered their own kingdom,
or part of the kingdomProtista.
A major research initiative to reconstruct the Red Algal Tree of Life (RedToL) using phylogenetic and genomic approaches is funded by the National Science Foundation as part of the Assembling the
Tree of Life Program.

Classification comparison

Classification system according to Classification system according to Pit

Orders Multicelluar? Example
Saunders and Hommersand 2004[28] Hwan Su Yoon et al. 2006[29] plugs?

Phylum Rhodophyta Wettstein

Subkingdom Rhodoplantae
Subphylum Cyanidiophytina
subphylum novus Cyanidioschyzon
Phylum Cyanidiophyta Cyanidiales No No
merolae
Class Cyanidiophyceae Merola et al. Class Cyanidiophyceae Merola
et al.

Phylum Rhodophyta Wettstein

Subphylum Rhodophytina
Subphylum Rhodellophytina subphylum novus
Rhodellales No No Rhodella
Class Rhodellophyceae Cavalier- Class Rhodellophyceae
Smith Cavalier-Smith

Subphylum Metarhodophytina Compsopogonales,

Class
Class Compsopogonophyceae Compsopogonophyceae Rhodochaetales, Yes No Compsopogon
Saunders et Hommersand Saunders et Hommersand Erythropeltidales

Class
Stylonematophyceae Rufusiales, Stylonematales Yes No Stylonema
classis nova

Subphylum Eurhodophytina Bangia,

Class Bangiophyceae Bangiales Yes Yes
Class Bangiophyceae Wettstein Wettstein "Porphyra"

Class
Porphyridiales Porphyridium
Porphyridiophyceae No No
cruentum
classis nova

Class Florideophyceae
Cronquist
Class Florideophyceae
Hildenbrandiales Yes Yes Hildenbrandia
Subclass Cronquist
Hildenbrandiophycidae

Batrachospermales,
Balliales, Balbianiales,
Subclass Nemaliales,
Yes Yes Nemalion
Nemaliophycidae Colaconematales,
Acrochaetiales,
Palmariales, Thoreales
Rhodogorgonales, Corallina
Yes Yes
Corallinales officinalis

Subclass
Ahnfeltiales, Pihiellales Yes Yes Ahnfeltia
Ahnfeltiophycidae

Bonnemaisoniales,
Gigartinales, Gelidiales,
Gracilariales,
Subclass
Halymeniales, Yes Yes Gelidium
Rhodymeniophycidae
Rhodymeniales,
Nemastomatales,
Plocamiales, Ceramiales

Some sources (such as Lee) place all red algae into the class "Rhodophyceae". (Lee's ganization
or [30] )
is not a comprehensive classification, but a selection of orders considered common or important.

A subphylum - Proteorhodophytina - has been proposed to encompass the existing classes Compsopogonophyceae, Porphyridiophyceae, Rhodellophyceae and Stylonematophyceae.[31] This proposal
was made on the basis of the analysis of the plastid genomes.

Species of red algae

[3] but the taxonomy is in constant flux with new species described each year
Over 7,000 species are currently described for the red algae, .[28][29] The vast majority of these are marine with about 200 that
live only in fresh water.

Some examples of species and genera of red algae are:

Cyanidioschyzon merolae, a primitive red alga

Atractophora hypnoides
Gelidiella calcicola
 Lemanea, a freshwater genus
Palmaria palmata, dulse
Schmitzia hiscockiana
Chondrus crispus, Irish moss
Mastocarpus stellatus
Vanvoorstia bennettiana, became extinct in the early 20th century
Acrochaetium efflorescens
Audouinella, with freshwater as well as marine species
Polysiphonia ceramiaeformis, banded siphon weed

Morphology
Red algae have double cell walls.[32] The outer layers contain the polysaccharides agarose and agaropectin that can be extracted from the cell walls by boiling as agar.[32] The internal walls are mostly
cellulose.[32]

Pit connections and pit plugs

Pit connections
Pit connections and pit plugs are unique and distinctive features of red algae that form during the process of cytokinesis following mitosis.[33][34] 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.

Shortly after the pit connection is formed, cytoplasmic continuity is blocked by the generation of a pit plug, which is deposited in the wall gap that connects the cells.

Connections between cells having a common parent cell are called primary pit connections. Because apical growth is the norm in red algae, most cells have two primary pit 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
Ceramiales.[34]
cell that then fuses to an adjacent cell. Patterns of secondary pit connections can be seen in the order

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 fofthe plug.

Function
[35]
The pit connections have been suggested to function as structural reinforcement, or as avenues for cell-to-cell communication and transport in red algae, however little data supports this hypothesis.

Reproduction
[2]
The reproductive cycle of red algae may be triggered by factors such as day length.

Fertilization
Red algae lack motile sperm. Hence, they rely on water currents to transport theirgametes to the female organs – although their sperm are capable of "gliding" to acarpogonium'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]

Upon their collision, the walls of the spermatium and carpogonium dissolve. The male nucleus divides and moves into the carpogonium; one half of the nucleus merges with the carpogonium's
nucleus.[2]

[2]
The polyamine spermine is produced, which triggers carpospore production.

[2]
Spermatangia may have long, delicate appendages, which increase their chances of "hooking up".

Life cycle
They display alternation of generations; in addition to gametophyte generation, many have two sporophyte generations, the carposporophyte-producing carpospores, which germinate into a
[2] The gametophyte is typically (but not always) identical to the tetrasporophyte.
tetrasporophyte – this produces spore tetrads, which dissociate and germinate into gametophytes. [36]

Carpospores may also germinate directly into thalloid gametophytes, or the carposporophytes may produce a tetraspore without going through a (free-living) tetrasporophyte phase.[36] Tetrasporangia
[2]
may be arranged in a row (zonate), in a cross (cruciate), or in a tetrad.

The carposporophyte may be enclosed within the gametophyte, which may cover it with branches to formcystocarp.
a [36]

These case studies may be helpful to understand some of the life histories algae may display:

In a simple case, such asRhodochorton investiens:

In the Carposporophyte: a spermatium merges with a trichogyne (a long hair on the female sexual or
gan), which then divides to form carposporangia – which produce carpospores.

Carpospores germinate into gametophytes, which produce sporophytes. Both of these are very similar; they produce monospores from monosporangia "just below a cross wall in a filament"[2] and their
[2]
spores are "liberated through apex of sporangial cell."

The spores of a sporophyte produce either tetrasporophytes. Monospores produced by this phase germinate immediately, with no resting phase, to form an identical copy of parent. Tetrasporophytes may
[2]
also produce a carpospore, which germinates to form another tetrasporophyte.

[2]
The gametophyte may replicate using monospores, but produces sperm in spermatangia, and "eggs"(?) in carpogonium.
A rather different example is Porphyra gardneri:

In its diploid phase, a carpospore can germinate to form a filamentous "conchocelis stage", which can also self-replicate using monospores. The conchocelis stage eventually produces conchosporangia.
The resulting conchospore germinates to form a tiny prothallus with rhizoids, which develops to a cm-scale leafy thallus. This too can reproduce via monospores, which are produced inside the thallus
itself.[2] They can also reproduce via spermatia, produced internally
, which are released to meet a prospective carpogonium in itsconceptacle.[2]

Chemistry
The δ13C values of red algae reflect their lifestyles. The largest difference results from their photosynthetic metabolic pathway: algae that use HCO3 as a
Algal group δ13C range[37]
carbon source have less negativeδ13C values than those that only use CO2.[37] An additional difference of about 1.71‰ separates groups intertidal from
those below the lowest tide line, which are never exposed to atmospheric carbon. The latter group uses the more 13C-negative CO2 dissolved in sea HCO3-using red −22.5‰ to
water, whereas those with access to atmospheric carbon reflect the more positive signature of thiseserve.
r algae −9.6‰
CO2-using red −34.5‰ to
Photosynthetic pigments of Rhodophyta are chlorophylls a and d. Red algae are red due to phycoerythrin. They contain the sulfated polysaccharide
algae −29.9‰
carrageenan in the amorphous sections of their cell walls, although red algae from the genus Porphyra contain porphyran. They also produce a specific
type of tannin called phlorotannins, but in lower amount than brown algae do. Brown algae −20.8‰ to
−10.5‰
Genomes of red algae Green algae −20.3‰ to
−8.8‰
Complete genome sequences are only available for 6 species of red algae, including 4 published in 2013.

Cyanidioschyzon merolae, Cyanidiophyceae[38][39]

Galdieria sulphuraria, Cyanidiophyceae[40]
Pyropia yezoensis, Bangiophyceae[41]
Chondrus crispus, Florideophyceae[42]
Porphyridium purpureum, Porphyridiophyceae[43]
Porphyra umbilicalis, Bangiophyceae[44]

Fossil record
One of the oldest fossils identified as a red alga is also the oldest fossil eukaryote that belongs to a specific modern taxon. Bangiomorpha pubescens, a multicellular fossil from arctic Canada, strongly
[1]
resembles the modern red algaBangia despite occurring in rocks dating to 1.2 billion years ago.

Two kinds of fossils resembling red algae were found sometime between 2006 and 2011 in well-preserved sedimentary rocks in Chitrakoot, central India. The presumed red algae lie embedded in fossil
mats of cyanobacteria, called stromatolites, in 1.6 billion-year [45]
-old Indian phosphorite — making them the oldest plant-like fossils ever found by about 400 million years.

Red algae are important builders oflimestone reefs. The earliest such coralline algae, thesolenopores, are known from the Cambrian period. Other algae of different origins filled 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.[46] Thallophytes resembling coralline red algae are known from the late Proterozoic
Doushantuo formation.[47]

Relationship to other algae

Chromista and Alveolata algae (e.g., chrysophytes, diatoms, phaeophytes, dinophytes) seem to have evolved from bikonts that have acquired red algae as endosymbionts. According to this theory, over
endosymbiotic theory is supported by various structural andgenetic similarities.[48]
time these endosymbiont red algae have evolved to become chloroplasts. This part of

Human consumption
Several species are important food crops, in particular members of the genus Porphyra, variously known as nori (Japan), gim (Korea), or laver (Britain). Dulse (Palmaria palmata)[49] is another
important British species.[50] These rhodophyte foods are high in vitamins and protein and are easily grown; for example,
nori cultivation in Japan goes back more than three centuries.

In East and Southeast Asia,agar is most commonly produced fromGelidium amansii.

Gallery
 Cyanidium sp. Porphyra sp., haploid Chondrus crispus Gracilaria sp. Corallina officinalis sp. Laurencia sp.
(Cyanidiophyceae) and diploid (Florideophyceae: (Florideophyceae: (Florideophyceae: (Florideophyceae:
(Bangiophyceae) Gigartinales) Gracilariales) Corallinales) Ceramiales)

Some red algae are

iridescent when not
covered with water

See also
Brown algae
Green algae
History of phycology

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