Temporal range: Mesoproterozoic–present
Classification is currently disputed. See Taxonomy.
The red algae, or Rhodophyta (// roh-DOF-fit-tə or // ROH-də-FY-tə; from Ancient Greek: ῥόδον rhodon, "rose" and φυτόν phyton, "plant"), are one of the oldest groups of eukaryotic algae. The Rhodophyta also contains one of the largest phyla of algae, containing over 7,000 currently recognized species with taxonomic revisions ongoing. The majority of species (6,793) are found in the Florideophyceae (class), and consist of mostly multicellular, marine algae, including many notable seaweeds. Although most species are marine, approximately 5% of the red algae occur in freshwater environments.
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. Red algae store sugars as floridean starch, which is a type of starch that consists of highly branched amylopectin without amylose, 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 generations that may have three generations rather than two.
Chloroplasts evolved following an endosymbiotic event between an ancestral, photosynthetic cyanobacterium and an early eukarytoic Phagotroph. 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. In the red algae, a secondary endosymbiosis event involving an ancestral red alga and a heterotrophic eukaryote resulted in the evolution and diversification of several other photosynthetic lineages.
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.
- 1 Habitat
- 2 Fossil record
- 3 Taxonomy
- 4 Species of red algae
- 5 Genomes of red algae
- 6 Relationship to Chromalveolata chloroplasts
- 7 Chemistry
- 8 Morphology
- 9 Pit connections and pit plugs
- 10 Reproduction
- 11 Human consumption
- 12 See also
- 13 References
- 14 External links
Unicellular members of the Cyanidiophyceae are thermoacidophiles and are found in sulphuric hot springs and other acidic environments. The remaining taxa are found in marine and freshwater environments. Most rhodophytes are marine with a worldwide distribution, and are often found at greater depths compared to other seaweeds because of dominance in certain pigments (i.e., Phycoerythrin) within their chloroplasts. Some marine species are found on sandy shores, while most others can be found attached to rocky substrata. Freshwater species account for 5% of red algal diversity, but they also have a worldwide distribution in various habitats; they generally prefer clean, high-flow streams with clear waters and rocky bottoms, but with some exceptions. A few freshwater species are found in black waters with sandy bottoms  and even fewer are found in more lentic waters. Both marine and freshwater taxa are represented by free-living macroalgal forms and smaller endo/epiphytic/zoic forms, meaning they live in or on other algae, plants, and animals  In addition, some marine species have adopted a parasitic lifestyle and may be found on closely or more distantly related red algal hosts 
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 resembles the modern red alga Bangia despite occurring in rocks dating to 1.2 billion years ago.
Red algae are important builders of limestone reefs. The earliest such coralline algae, the solenopores, 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. Thallophytes resembling coralline red algae are known from the late Proterozoic Doushantuo formation.
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."
Many studies published since Adl et al. 2005 have provided evidence that is in agreement for monophyly in the Archaeplastida (including red algae). However, other studies have suggested Archaeplastida is paraphyletic. As of January 2011[update], 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 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 kingdom Protista.
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.
Some sources (such as Lee) place all red algae into the class "Rhodophyceae". (Lee's organization is not a comprehensive classification, but a selection of orders considered common or important.)
Species of red algae
Over 7,000 species are currently described for the red algae, but the taxonomy is in constant flux with new species described each year. 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
Genomes of red algae
Complete genome sequences are only available for 5 species of red algae, including 4 published in 2013.
- Cyanidioschyzon merolae, Cyanidiophyceae
- Galdieria sulphuraria, Cyanidiophyceae
- Pyropia yezoensis, Bangiophyceae
- Chondrus crispus, Florideophyceae
- Porphyridium purpureum, Porphyridiophyceae
Relationship to Chromalveolata chloroplasts
Chromalveolates seem to have evolved from bikonts that have acquired red algae as endosymbionts. According to this theory, over time these bikonts and their endosymbiont red algae have evolved to become chromalveolates and their chloroplasts. This part of endosymbiotic theory is supported by various structural and genetic similarities.
|Algal group||δ13C range|
|HCO3-using red algae||−22.5‰ to −9.6‰|
|CO2-using red algae||−34.5‰ to −29.9‰|
|Brown algae||−20.8‰ to −10.5‰|
|Green algae||−20.3‰ to −8.8‰|
The δ13C values of red algae reflect their lifestyles. The largest difference results from their photosynthetic metabolic pathway: algae that use HCO3 as a carbon source have far more negative δ13C values than those that only use CO2. 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 water, whereas those with access to atmospheric carbon reflect the more positive signature of this reserve.
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 specific type of tannin called phlorotannins, but in lower amount than brown algae do.
Red algae have double cell walls. The outer layers contain the polysaccharides agarose and agaropectin that can be extracted from the cell walls by boiling as agar. The internal walls are mostly cellulose.
Pit connections and pit plugs
Pit connections and pit plugs are unique and distinctive features of red algae that form during the process of cytokinesis following mitosis. 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 cell that then fuses to an adjacent cell. Patterns of secondary pit connections can be seen in the order Ceramiales.
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 off the plug.
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.
The reproductive cycle of red algae may be triggered by factors such as day length.
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.
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.
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. The gametophyte is typically (but not always) identical to the tetrasporophyte.
Carpospores may also germinate directly into thalloid gametophytes, or the carposporophytes may produce a tetraspore without going through a (free-living) tetrasporophyte phase. Tetrasporangia 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 form a cystocarp.
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 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" and their 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 also produce a carpospore, which germinates to form another tetrasporophyte.[verification needed]
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. They can also reproduce via spermatia, produced internally, which are released to meet a prospective carpogonium in its conceptacle.
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) is another important British species. 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 from Gelidium amansii.
- N. J. Butterfield (2000). "Bangiomorpha pubescens n. gen., n. sp.: implications for the evolution of sex, multicellularity, and the Mesoproterozoic/Neoproterozoic radiation of eukaryotes". Paleobiology. 26 (3): 386–404. doi:10.1666/0094-8373(2000)026<0386:BPNGNS>2.0.CO;2. ISSN 0094-8373.
- Lee, R.E. (2008). Phycology, 4th edition. Cambridge University Press. ISBN 978-0-521-63883-8.
- Guiry, M.D.; Guiry, G.M. (2016). "Algaebase". www.algaebase.org. Retrieved November 20, 2016.
- D. Thomas (2002). Seaweeds. Life Series. Natural History Museum, London. ISBN 0-565-09175-1.
- Sheath, Robert G. (1984). "The biology of freshwater red algae". Progress Phycological Research. 3: 89–157.
- W. J. Woelkerling (1990). "An introduction". In K. M. Cole; R. G. Sheath. Biology of the Red Algae. Cambridge University Press, Cambridge. pp. 1–6. ISBN 0-521-34301-1.
- Viola, R.; Nyvall, P.; Pedersén, M. (2001). "The unique features of starch metabolism in red algae.". Proceedings of the Royal Society of London B. 268: 1417=1422. doi:10.1098/rspb.2001.1644.
- Gould, S.B.; Waller, R.F.; McFadden, G.I. (2008). "Plastid Evolution". Annual Reviews in Plant Biology. 59: 491–517.
- McFadden, G.I. (2001). "Primary and Secondary Endosymbiosis and the Evolution of Plastids". Journal of Phycology. 37: 951–959.
- M. D. Guiry. "Rhodophyta: red algae". National University of Ireland, Galway. Archived from the original on 2007-05-04. Retrieved 2007-06-28.
- Ciniglia, C.; Yoon, H.; Pollio, A.; Bhattacharya, D. (2004). "Hidden biodiversity of the extremophilic Cyanidiales red algae". Molecular Ecology. 13: 1827–1838. doi:10.1111/j.1365-294X.2004.02180.x.
- Kain, J.M.; Norton, T.A. (1990). "Marine Ecology". In Cole, J.M.; Sheath, R.G. Biology of the Red Algae. Cambridge, U.K.: Cambridge University Press. pp. 377–423.
- Eloranta, P.; Kwandrans, J. (2004). "Indicator value of freshwater red algae in running waters for water quality assessment" (PDF). International Journal of Oceanography and Hydrobiology. XXXIII (1): 47–54. ISSN 1730-413X.
- Vis, M.L.; Sheath, R.G.; Chiasson, W.B. (2008). "A survey of Rhodophyta and associated macroalgae from coastal streams in French Guiana". Cryptogamie Algologie. 25: 161–174.
- Sheath, R.G.; Hambrook, J.A. (1990). "Freshwater Ecology". In Cole, K.M.; Sheath, R.G. Biology of the Red Algae. Cambridge, U.K.: Cambridge University Press. pp. 423–453.
- Goff, L.J. (1982). "The biology of parasitic red algae". Progress Phycological Research. 1: 289–369.
- Salomaki, E.D.; Lane, C.E. (2014). "Are all red algal parasites cut from the same cloth?". Acta Societatis Botanicorum Poloniae. 83: 369–375. doi:10.5586/asbp.2014.047.
- Grant, S. W. F.; Knoll, A. H.; Germs, G. J. B. (1991). "Probable Calcified Metaphytes in the Latest Proterozoic Nama Group, Namibia: Origin, Diagenesis, and Implications". Journal of Paleontology. JSTOR. 65 (1): 1–18. JSTOR 1305691. PMID 11538648.
- Yun, Z.; Xun-lal, Y. (1992). "New data on multicellular thallophytes and fragments of cellular tissues from Late Proterozoic phosphate rocks, South China". Lethaia. 25 (1): 1–18. doi:10.1111/j.1502-3931.1992.tb01788.x.
- Adl, Sina M.; et al. (2005). "The New Higher Level Classification of Eukaryotes with Emphasis on the Taxonomy of Protists". Journal of Eukaryotic Microbiology. 52 (5): 399–451. doi:10.1111/j.1550-7408.2005.00053.x. PMID 16248873
- Fabien Burki, Kamran Shalchian-Tabrizi, Marianne Minge, Åsmund Skjæveland, Sergey I. Nikolaev, Kjetill S. Jakobsen, Jan Pawlowski (2007). Butler, Geraldine, ed. "Phylogenomics Reshuffles the Eukaryotic Supergroups". PLoS ONE. 2 (8): e790. doi:10.1371/journal.pone.0000790. PMC . PMID 17726520.
- Burki, Fabien; Inagaki, Yuji; Bråte, Jon; Archibald, John M.; Keeling, Patrick J.; Cavalier-Smith, Thomas; Sakaguchi, Miako; Hashimoto, Tetsuo; Horak, Ales; Kumar, Surendra; Klaveness, Dag; Jakobsen, Kjetill S.; Pawlowski, Jan; Shalchian-Tabrizi, Kamran (2009). "Large-Scale Phylogenomic Analyses Reveal That Two Enigmatic Protist Lineages, Telonemia and Centroheliozoa, Are Related to Photosynthetic Chromalveolates". Genome Biology and Evolution. 1: 231–8. doi:10.1093/gbe/evp022. PMC . PMID 20333193.
- Cavalier-Smith, Thomas (2009). "Kingdoms Protozoa and Chromista and the eozoan root of the eukaryotic tree". Biology Letters. 6 (3): 342–5. doi:10.1098/rsbl.2009.0948. PMC . PMID 20031978.
- Rogozin, I.B.; Basu, M.K.; Csürös, M. & Koonin, E.V. (2009). "Analysis of Rare Genomic Changes Does Not Support the Unikont–Bikont Phylogeny and Suggests Cyanobacterial Symbiosis as the Point of Primary Radiation of Eukaryotes". Genome Biology and Evolution. 1: 99–113. doi:10.1093/gbe/evp011. PMC . PMID 20333181.
- Kim, E.; Graham, L.E. & Graham, Linda E. (2008). Redfield, Rosemary Jeanne, ed. "EEF2 analysis challenges the monophyly of Archaeplastida and Chromalveolata". PLoS ONE. 3 (7): e2621. doi:10.1371/journal.pone.0002621. PMC . PMID 18612431
- Nozaki, H.; Maruyama, S.; Matsuzaki, M.; Nakada, T.; Kato, S.; Misawa, K. (2009). "Phylogenetic positions of Glaucophyta, green plants (Archaeplastida) and Haptophyta (Chromalveolata) as deduced from slowly evolving nuclear genes". Molecular Phylogenetics and Evolution. 53 (3): 872–880. doi:10.1016/j.ympev.2009.08.015. PMID 19698794
- G. W. Saunders & M. H. Hommersand (2004). "Assessing red algal supraordinal diversity and taxonomy in the context of contemporary systematic data". American Journal of Botany. 91 (10): 1494–1507. doi:10.3732/ajb.91.10.1494.
- Hwan Su Yoon, K. M. Müller, R. G. Sheath, F. D. Ott & D. Bhattacharya (2006). "Defining the major lineages of red algae (Rhodophyta)" (PDF). Journal of Phycology. 42 (2): 482–492. doi:10.1111/j.1529-8817.2006.00210.x.
- Robert Edward Lee (2008). Phycology. Cambridge University Press. p. 107. ISBN 978-0-521-68277-0. Retrieved 31 January 2011.
- Matsuzaki; et al. (April 2004). "(April 8, 2004) Genome sequence of the ultrasmall unicellular red alga Cyanidioschyzon merolae 10D". Nature. 428 (6983): 653–7. doi:10.1038/nature02398. PMID 15071595.
- Nozaki et al. (July 10, 2007) A 100%-complete sequence reveals unusually simple genomic features in the hot-spring red alga Cyanidioschyzon merolae. BMC Biol. 2007; 5: 28. doi:10.1186/1741-7007-5-28
- Schönknecht; et al. (Mar 2013). "(March 8, 2013) Gene Transfer from Bacteria and Archaea Facilitated Evolution of an Extremophilic Eukaryote". Science. 339 (6124): 1207–1210. doi:10.1126/science.1231707. PMID 23471408.
- Nakamura; et al. "(March 11, 2013) The First Symbiont-Free Genome Sequence of Marine Red Alga, Susabi-nori (Pyropia yezoensis)". PLoS ONE. 8 (3): e57122. doi:10.1371/journal.pone.0057122.
- Collen; et al. (2013). "Genome structure and metabolic features in the red seaweed Chondrus crispus shed light on evolution of the Archaeplastida". PNAS. 110: 5247–5252. doi:10.1073/pnas.1221259110.
- Bhattacharya; et al. (2013). "Genome of the red alga Porphyridium purpureum". Nature Communications. 4: 1941. doi:10.1038/ncomms2931.
- Summarised in Cavalier-Smith, Thomas (April 2000). "Membrane heredity and early chloroplast evolution". Trends in Plant Science. 5 (4): 174–182. doi:10.1016/S1360-1385(00)01598-3. PMID 10740299.
- Maberly, S. C.; Raven, J. A.; Johnston, A. M. (1992). "Discrimination between 12C and 13C by marine plants". Oecologia. 91 (4): 481. doi:10.1007/BF00650320. JSTOR 4220100.
- Fritsch, F. E. (1945), The structure and reproduction of the algae, Cambridge: Cambridge Univ. Press, ISBN 0521050421, OCLC 223742770
- Clinton JD, Scott FM, Bowler E (November–December 1961). "A Light- and Electron-Microscopic Survey of Algal Cell Walls. I. Phaeophyta and Rhodophyta". American Journal of Botany. Botanical Society of America. 48 (10): 925–934. doi:10.2307/2439535. JSTOR 2439535.
- Lee RE (2008). Phycology (4th ed.). Cambridge University Press. ISBN 978-0-521-63883-8.
- "Pit Plugs". FHL Marine Botany. Retrieved 2016-06-30.
- Kohlmeyer, J. (February 1975). "New Clues to the Possible Origin of Ascomycetes". BioScience. American Institute of Biological Sciences. 25 (2): 86–93. doi:10.2307/1297108. JSTOR 1297108.
- "Dulse: Palmaria palmata". Quality Sea Veg. Retrieved 2007-06-28.
- T. F. Mumford & A. Muira (1988). "Porphyra as food: cultivation and economics". In C. A. Lembi & J. Waaland. Algae and Human Affairs. Cambridge University Press, Cambridge. ISBN 0-521-32115-8.