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Algae
Laurencia, a marine genus of Red Algae from Hawaii.
Laurencia, a marine genus of Red Algae from Hawaii.
Scientific classification
Domain: Eukaryota
Included groups
Excluded groups
The lineage of algae according to Thomas Cavalier-Smith. The exact number and placement of endosymbiotic events is not yet clear, so this diagram can be taken only as a general guide[1][2] It represents the most parsimonious way of explaining the three types of endosymbiotic origins of chloroplasts. These types include the endosymbiotic events of cyanobacteria, red algae and green algae, leading to the hypothesis of the supergroups Archaeplastida, Chromalveolata and Cabozoa respectively. However, the monophyly of Cabozoa has been refuted and the monophylies of Archaeplastida and Chromalveolata are currently strongly challenged. Endosymbiotic events are noted by dotted lines.
Algae (pronounced /ˈældʒiː/ or /ˈælɡiː/; singular alga /ˈælɡə/, Latin for "seaweed") are a large and diverse group of simple, typically autotrophic organisms, ranging from unicellular to multicellular forms. The largest and most complex marine forms are called seaweeds. They are photosynthetic, like plants, and "simple" because they lack the many distinct organs found in land plants.
Though the prokaryotic Cyanobacteria (commonly referred to as blue-green algae) were traditionally included as "algae" in older textbooks, many modern sources regard this as outdated[3] as they are now considered to be closely related to bacteria.[4] The term algae is now restricted to eukaryotic organisms.[5] All true algae therefore have a nucleus enclosed within a membrane and chloroplasts bound in one or more membranes.[3][6] Algae constitute a paraphyletic and polyphyletic group,[3] as they do not include all the descendants of the last universal ancestor nor do they all descend from a common algal ancestor, although their chloroplasts seem to have a single origin.[1] Diatoms are also examples of algae.
Algae lack the various structures that characterize land plants, such as phyllids (leaves) and rhizoids in nonvascular plants, or leaves, roots, and other organs that are found in tracheophytes (vascular plants). Many are photoautotrophic, although some groups contain members that are mixotrophic, deriving energy both from photosynthesis and uptake of organic carbon either by osmotrophy, myzotrophy, or phagotrophy. Some unicellular species rely entirely on external energy sources and have limited or no photosynthetic apparatus.
Nearly all algae have photosynthetic machinery ultimately derived from the Cyanobacteria, and so produce oxygen as a by-product of photosynthesis, unlike other photosynthetic bacteria such as purple and green sulfur bacteria. Fossilized filamentous algae from the Vindhya basin have been dated back to 1.6 to 1.7 billion years ago.[7]
The first alga to have its genome sequenced was Cyanidioschyzon merolae.

Contents

Etymology and study

Title page of Samuel Gottlieb Gmelin, Historia Fucorum, dated 1768.
The singular alga is the Latin word for a particular seaweed and retains that meaning in English.[8] The etymology is obscure. Although some speculate that it is related to Latin algēre, "be cold",[9] there is no known reason to associate seaweed with temperature. A more likely source is alliga, "binding, entwining."[10] Since Algae has become a biological classification, alga can also mean one classification under Algae, parallel to a fungus being a species of fungi, a plant being a species of plant, and so on.
The ancient Greek word for seaweed was φῦκος (fūkos or phykos), which could mean either the seaweed, probably Red Algae, or a red dye derived from it. The Latinization, fūcus, meant primarily the cosmetic rouge. The etymology is uncertain, but a strong candidate has long been some word related to the Biblical פוך (pūk), "paint" (if not that word itself), a cosmetic eye-shadow used by the ancient Egyptians and other inhabitants of the eastern Mediterranean. It could be any color: black, red, green, blue.[11]
Accordingly the modern study of marine and freshwater algae is called either phycology or algology. The name Fucus appears in a number of taxa.

Classification

False-colour Scanning electron micrograph of the unicellular coccolithophore, Gephyrocapsa oceanica.
While Cyanobacteria have been traditionally included among the Algae, recent works usually exclude them due to large differences such as the lack of membrane-bound organelles, the presence of a single circular chromosome, the presence of peptidoglycan in the cell walls, and ribosomes different in size and content from those of the Eukaryotes.[12][13] Rather than in chloroplasts, they conduct photosynthesis on specialized infolded cytoplasmic membranes called thylakoid membranes. Therefore, they differ significantly from the Algae despite occupying similar ecological niches.
By modern definitions Algae are Eukaryotes and conduct photosynthesis within membrane-bound organelles called chloroplasts. Chloroplasts contain circular DNA and are similar in structure to Cyanobacteria, presumably representing reduced cyanobacterial endosymbionts. The exact nature of the chloroplasts is different among the different lines of Algae, reflecting different endosymbiotic events. The table below describes the composition of the three major groups of Algae. Their lineage relationships are shown in the figure in the upper right. Many of these groups contain some members that are no longer photosynthetic. Some retain plastids, but not chloroplasts, while others have lost plastids entirely.
Phylogeny based on plastid.[14] not nucleocytoplasmic genealogy:

Cyanobacteria





Rhodoplasts











Chloroplasts



















Supergroup affiliation Members Endosymbiont Summary
Primoplantae/
Archaeplastida
Cyanobacteria These Algae have primary chloroplasts, i.e. the chloroplasts are surrounded by two membranes and probably developed through a single endosymbiotic event. The chloroplasts of Red Algae have chlorophylls a and c (often), and phycobilins, while those of Green Algae have chloroplasts with chlorophyll a and b. Higher plants are pigmented similarly to Green Algae and probably developed from them, and thus Chlorophyta is a sister taxon to the plants; sometimes they are grouped as Viridiplantae.
Excavata and Rhizaria Green Algae
These groups have green chloroplasts containing chlorophylls a and b [12]. Their chloroplasts are surrounded by four and three membranes, respectively, and were probably retained from ingested Green Algae.
Chlorarachniophytes, which belong to the phylum Cercozoa, contain a small nucleomorph, which is a relict of the algae's nucleus.
Euglenids, which belong to the phylum Euglenozoa, live primarily in freshwater and have chloroplasts with only three membranes. It has been suggested that the endosymbiotic Green Algae were acquired through myzocytosis rather than phagocytosis.
Chromista and Alveolata Red Algae
These groups have chloroplasts containing chlorophylls a and d, and phycobilins. The latter chlorophyll type is not known from any prokaryotes or primary chloroplasts, but genetic similarities with the Red Algae suggest a relationship there.
In the first three of these groups (Chromista), the chloroplast has four membranes, retaining a nucleomorph in Cryptomonads, and they likely share a common pigmented ancestor, although other evidence casts doubt on whether the Heterokonts, Haptophyta, and Cryptomonads are in fact more closely related to each other than to other groups.[2][15]
The typical dinoflagellate chloroplast has three membranes, but there is considerable diversity in chloroplasts within the group, and it appears there were a number of endosymbiotic events.[1] The Apicomplexa, a group of closely related parasites, also have plastids called apicoplasts. Apicoplasts are not photosynthetic but appear to have a common origin with Dinoflagellate chloroplasts.[1]
W.H.Harvey (1811—1866) was the first to divide the Algae into four divisions based on their pigmentation. This is the first use of a biochemical criterion in plant systematics. Harvey's four divisions are: Red Algae (Rhodophyta), Brown Algae (Heteromontophyta), Green Algae (Chlorophyta) and Diatomaceae.[16]

Relationship to higher plants

The first plants on earth evolved from shallow freshwater algae much like Chara some 400 million years ago. These probably had an isomorphic alternation of generations and were probably filamentous. Fossils of isolated land plant spores suggest land plants may have been around as long as 475 million years ago.[17][18]

Morphology

The kelp forest exhibit at the Monterey Bay Aquarium. A three-dimensional, multicellular thallus.
A range of algal morphologies are exhibited, and convergence of features in unrelated groups is common. The only groups to exhibit three dimensional multicellular thalli are the reds and browns, and some chlorophytes.[19] Apical growth is constrained to subsets of these groups: the florideophyte reds, various browns, and the charophytes.[19] The form of charophytes is quite different to those of reds and browns, because have distinct nodes, separated by internode 'stems'; whorls of branches reminiscent of the horsetails occur at the nodes.[19] Conceptacles are another polyphyletic trait; they appear in the coralline algae and the Hildenbrandiales, as well as the browns.[19]
Most of the simpler algae are unicellular flagellates or amoeboids, but colonial and non-motile forms have developed independently among several of the groups. Some of the more common organizational levels, more than one of which may occur in the life cycle of a species, are
  • Colonial: small, regular groups of motile cells
  • Capsoid: individual non-motile cells embedded in mucilage
  • Coccoid: individual non-motile cells with cell walls
  • Palmelloid: non-motile cells embedded in mucilage
  • Filamentous: a string of non-motile cells connected together, sometimes branching
  • Parenchymatous: cells forming a thallus with partial differentiation of tissues
In three lines even higher levels of organization have been reached, with full tissue differentiation. These are the brown algae,[20]—some of which may reach 50 m in length (kelps)[21]—the red algae,[22] and the green algae.[23] The most complex forms are found among the green algae (see Charales and Charophyta), in a lineage that eventually led to the higher land plants. The point where these non-algal plants begin and algae stop is usually taken to be the presence of reproductive organs with protective cell layers, a characteristic not found in the other alga groups.

Symbiotic algae

Some species of algae form symbiotic relationships with other organisms. In these symbioses, the algae supply photosynthates (organic substances) to the host organism providing protection to the algal cells. The host organism derives some or all of its energy requirements from the algae. Examples are as follows.

Lichens

Rock lichens in Ireland.
Lichens are defined by the International Association for Lichenology to be "an association of a fungus and a photosynthetic symbiont resulting in a stable vegetative body having a specific structure."[24] The fungi, or mycobionts, are from the Ascomycota with a few from the Basidiomycota. They are not found alone in nature but when they began to associate is not known.[25] One mycobiont associates with the same phycobiont species, rarely two, from the Green Algae, except that alternatively the mycobiont may associate with the same species of Cyanobacteria (hence "photobiont" is the more accurate term). A photobiont may be associated with many specific mycobionts or live independently; accordingly, lichens are named and classified as fungal species.[26] The association is termed a morphogenesis because the lichen has a form and capabilities not possessed by the symbiont species alone (they can be experimentally isolated). It is possible that the photobiont triggers otherwise latent genes in the mycobiont.[27]

Coral reefs

Floridian coral reef
Coral reefs are accumulated from the calcareous exoskeletons of marine invertebrates of the Scleractinia order; i.e., the Stony Corals. As animals they metabolize sugar and oxygen to obtain energy for their cell-building processes, including secretion of the exoskeleton, with water and carbon dioxide as byproducts. As the reef is the result of a favorable equilibrium between construction by the corals and destruction by marine erosion, the rate at which metabolism can proceed determines the growth or deterioration of the reef.
Algae of the Dinoflagellate phylum are often endosymbionts in the cells of marine invertebrates, where they accelerate host-cell metabolism by generating immediately available sugar and oxygen through photosynthesis using incident light and the carbon dioxide produced in the host. Endosymbiont algae in the Stony Corals are described by the term zooxanthellae, with the host Stony Corals called on that account hermatypic corals, which although not a taxon are not in healthy condition without their endosymbionts. Zooxanthellae belong almost entirely to the genus Symbiodinium.[28] The loss of Symbiodinium from the host is known as coral bleaching, a condition which unless corrected leads to the deterioration and loss of the reef.

Sea sponges

Green Algae live close to the surface of some sponges, for example, breadcrumb sponge (Halichondria panicea). The alga is thus protected from predators; the sponge is provided with oxygen and sugars which can account for 50 to 80% of sponge growth in some species.[29]

Life-cycle

Rhodophyta, Chlorophyta and Heterokontophyta, the three main algal Phyla, have life-cycles which show tremendous variation with considerable complexity. In general there is an asexual phase where the seaweed's cells are diploid, a sexual phase where the cells are haploid followed by fusion of the male and female gametes. Asexual reproduction is advantageous in that it permits efficient population increases, but less variation is possible. Sexual reproduction allows more variation, but is more costly. Often there is no strict alternation between the sporophyte and also because there is often an asexual phase, which could include the fragmentation of the thallus.[21][30][31]

Numbers

The Algal Collection of the U.S. National Herbarium (located in the National Museum of Natural History) consists of approximately 320500 dried specimens, which, although not exhaustive (no exhaustive collection exists), gives an idea of the order of magnitude of the number of algal species (that number remains unknown).[32] Estimates vary widely. For example, according to one standard textbook,[33] in the British Isles the UK Biodiversity Steering Group Report estimated there to be 20000 algal species in the UK. Another checklist reports only about 5000 species. Regarding the difference of about 15000 species, the text concludes: "It will require many detailed field surveys before it is possible to provide a reliable estimate of the total number of species ...."
Regional and group estimates have been made as well: 5000—5500 species of Red Algae worldwide, "some 1300 in Australian Seas,"[34] 400 seaweed species for the western coastline of South Africa,[35] 669 marine species from California (U.S.A.),[36] 642 in the check-list of Britain and Ireland,[37] and so on, but lacking any scientific basis or reliable sources, these numbers have no more credibility than the British ones mentioned above. Most estimates also omit the microscopic Algae, such as the phytoplankta, entirely.

Distribution

The topic of distribution of algal species has been fairly well studied since the founding of phytogeography in the mid-19th century AD.[38] Algae spread mainly by the dispersal of spores analogously to the dispersal of Plantae by seeds and spores. Spores are everywhere in all parts of the Earth: the waters fresh and marine, the atmosphere, free-floating and in precipitation or mixed with dust, the humus and in other organisms, such as humans. Whether a spore is to grow into an organism depends on the combination of the species and the environmental conditions.
The spores of fresh-water Algae are dispersed mainly by running water and wind, as well as by living carriers.[39] The bodies of water into which they are transported are chemically selective. Marine spores are spread by currents. Ocean water is temperature selective, resulting in phytogeographic zones, regions and provinces.[40]
To some degree the distribution of Algae is subject to floristic discontinuities caused by geographical features, such as Antarctica, long distances of ocean or general land masses. It is therefore possible to identify species occurring by locality, such as "Pacific Algae" or "North Sea Algae". When they occur out of their localities, it is usually possible to hypothesize a transport mechanism, such as the hulls of ships. For example, Ulva reticulata and Ulva fasciata travelled from the mainland to Hawaii in this manner.
Mapping is possible for select species only: "there are many valid examples of confined distribution patterns."[41] For example, Clathromorphum is an arctic genus and is not mapped far south of there.[42] On the other hand, scientists regard the overall data as insufficient due to the "difficulties of undertaking such studies."[43]

Locations

Phytoplankton, Lake Chuzenji
Algae are prominent in bodies of water, common in terrestrial environments and are found in unusual environments, such as on snow and on ice. Seaweeds grow mostly in shallow marine waters, under 100 metres (330 ft); however some have been recorded to a depth of 360 metres (1,180 ft)[44]
The various sorts of algae play significant roles in aquatic ecology. Microscopic forms that live suspended in the water column (phytoplankton) provide the food base for most marine food chains. In very high densities (algal blooms) these algae may discolor the water and outcompete, poison, or asphyxiate other life forms.
Algae are variously sensitive to different factors, which has made them useful as biological indicators in the Ballantine Scale and its modification.

Uses

Harvesting Algae

Agar

Agar, an Algae derivative, has a number of commercial uses.[45]

Alginates

Between 100,000 and 170,000 wet tons of Macrocystis are harvested annually in California for alginate extraction and abalone feed.[46][47]

Energy source

To be competitive and independent from fluctuating support from (local) policy on the long run, biofuels should equal or beat the cost level of fossil fuels. Here, algae based fuels hold great promise, directly related to the potential to produce more biomass per unit area in a year than any other form of biomass. The break-even point for algae-based biofuels should be within reach in about ten years.[citation needed]

Fertilizer

Seaweed is used as a fertilizer.
For centuries seaweed has been used as a fertilizer; George Owen of Henllys writing in the 16th century referring to drift weed in South Wales:[48]
This kind of ore they often gather and lay on great heapes, where it heteth and rotteth, and will have a strong and loathsome smell; when being so rotten they cast on the land, as they do their muck, and thereof springeth good corn, especially barley ... After spring-tydes or great rigs of the sea, they fetch it in sacks on horse backes, and carie the same three, four, or five miles, and cast it on the lande, which doth very much better the ground for corn and grass.
Today Algae are used by humans in many ways; for example, as fertilizers, soil conditioners and livestock feed.[49] Aquatic and microscopic species are cultured in clear tanks or ponds and are either harvested or used to treat effluents pumped through the ponds. Algaculture on a large scale is an important type of aquaculture in some places. Maerl is commonly used as a soil conditioner.

Nutrition

Seaweed gardens on Inisheer.
Naturally growing seaweeds are an important source of food, especially in Asia. They provide many vitamins including: A, B1, B2, B6, niacin and C, and are rich in iodine, potassium, iron, magnesium and calcium.[50] In addition commercially cultivated microalgae, including both Algae and Cyanobacteria, are marketed as nutritional supplements, such as Spirulina,[51] Chlorella and the Vitamin-C supplement, Dunaliella, high in beta-carotene.
Algae are national foods of many nations: China consumes more than 70 species, including fat choy, a cyanobacterium considered a vegetable; Japan, over 20 species;[52] Ireland, dulse; Chile, cochayuyo.[53] Laver is used to make "laver bread" in Wales where it is known as bara lawr; in Korea, gim; in Japan, nori and aonori. It is also used along the west coast of North America from California to British Columbia, in Hawaii and by the Māori of New Zealand. Sea lettuce and badderlocks are a salad ingredient in Scotland, Ireland, Greenland and Iceland.
Dulse, a food.
The oils from some Algae have high levels of unsaturated fatty acids. For example, Parietochloris incisa is very high in arachidonic acid, where it reaches up to 47% of the triglyceride pool.[54] Some varieties of Algae favored by vegetarianism and veganism contain the long-chain, essential omega-3 fatty acids, Docosahexaenoic acid (DHA) and Eicosapentaenoic acid (EPA), in addition to vitamin B12.[citation needed] The vitamin B12 in algae is not biologically active. Fish oil contains the omega-3 fatty acids, but the original source is algae (microalgae in particular), which are eaten by marine life such as copepods and are passed up the food chain.[55] Algae has emerged in recent years as a popular source of omega-3 fatty acids for vegetarians who cannot get long-chain EPA and DHA from other vegetarian sources such as flaxseed oil, which only contains the short-chain Alpha-Linolenic acid (ALA).

Pollution control

  • Sewage can be treated with algae, reducing the need for greater amounts of toxic chemicals than are already used.
  • Algae can be used to capture fertilizers in runoff from farms. When subsequently harvested, the enriched algae itself can be used as fertilizer.

Pigments

The natural pigments produced by algae can be used as an alternative to chemical dyes and coloring agents.[56]

Stabilizing substances

Carrageenan, from the red alga Chondrus crispus, is used as a stabiliser in milk products.

See also

Notes

  1. ^ a b c d Patrick J. Keeling (2004). "Diversity and evolutionary history of plastids and their hosts". American Journal of Botany 91: 1481–1493. doi:10.3732/ajb.91.10.1481. http://www.amjbot.org/cgi/content/full/91/10/1481. 
  2. ^ a b Laura Wegener Parfrey, Erika Barbero, Elyse Lasser, Micah Dunthorn, Debashish Bhattacharya, David J Patterson, and Laura A Katz (December 2006). "Evaluating Support for the Current Classification of Eukaryotic Diversity". PLoS Genet. 2 (12): e220. doi:10.1371/journal.pgen.0020220. PMID 17194223. PMC 1713255. http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=1713255. 
  3. ^ a b c Nabors, Murray W. (2004). Introduction to Botany. San Francisco, CA: Pearson Education, Inc. ISBN 0805344160. 
  4. ^ Ed. Guiry, M.D., John, D.M., Rindi, F and McCarthy, T.K. 2007. New Survey of Clare Island Volume 6: The Freshwater and Terrestrial Algae. Royal Irish Academy. isbn 13: 978-1-904890-31-7
  5. ^ Allaby, M ed. (1992). "Algae". The Concise Dictionary of Botany. Oxford: Oxford University Press. 
  6. ^ Round (1981)
  7. ^ Bengtson, S.; Belivanova, V.; Rasmussen, B.; Whitehouse, M. (May 2009). "The controversial "Cambrian" fossils of the Vindhyan are real but more than a billion years older". Proceedings of the National Academy of Sciences of the United States of America 106 (19): 7729–7734. doi:10.1073/pnas.0812460106. ISSN 0027-8424. PMID 19416859.  edit
  8. ^ "alga, algae". Webster's Third New International Dictionary of the English Language Unabridged with Seven Language Dictionary. 1. Encyclopedia Britannica, Inc. 1986. 
  9. ^ Partridge, Eric (1983). "algae". Origins. 
  10. ^ Lewis, Charlton T.; Charles Short (1879). "alga". alga. Oxford: Clarendon Press. ISBN 0198642016. http://www.perseus.tufts.edu/cgi-bin/ptext?layout.reflang=la;layout.refdoc=Perseus%3Atext%3A1999.04.0059;layout.reflookup=Alga;layout.refcit=;doc=Perseus%3Atext%3A1999.04.0059%3Aentry%3D%231812. 
  11. ^ Cheyne, Thomas Kelly; John Sutherland Black (1899–1903). "Paint". Encyclopædia Biblica – A Dictionary of the Bible. 3. New York: Macmillan Co.. pp. 3524–3525.  Downloadable Google Books.
  12. ^ a b Losos, Jonathan B.; Mason, Kenneth A.; Singer, Susan R. (2007). Biology (8 ed.). McGraw-Hill. ISBN 0073041106. 
  13. ^ Jochem, Frank J. "Botany 4404 Lecture Notes". Florida International University (FIU). http://www.jochemnet.de/fiu/bot4404/BOT4404_12.html. Retrieved 2008-12-20. 
  14. ^ Bhattacharya, D.; Medlin, L. (1998). "Algal Phylogeny and the Origin of Land Plants". Plant Physiology 116 (1): 9–15. doi:10.1104/pp.116.1.9. http://www.plantphysiol.org/cgi/reprint/116/1/9.pdf. 
  15. ^ Burki F, Shalchian-Tabrizi K, Minge M, Skjæveland Å, Nikolaev SI, et al. (2007). "Phylogenomics Reshuffles the Eukaryotic Supergroups". PLoS ONE 2 (8: e790): e790. doi:10.1371/journal.pone.0000790. PMID 17726520. 
  16. ^ Dixon, P S (1973). Biology of the Rhodophyta. Edinburgh: Oliver & Boyd. p. 232. ISBN 005002485X. ISBN 005002485X. 
  17. ^ Ivan Noble (18 September 2003). "When plants conquered land". BBC. http://news.bbc.co.uk/1/hi/sci/tech/3117034.stm. 
  18. ^ Wellman, C.H.; Osterloff, P.L.; Mohiuddin, U. (2003). "Fragments of the earliest land plants". Nature 425 (6955): 282–285. doi:10.1038/nature01884. PMID 13679913. 
  19. ^ a b c d Xiao, S.; Knoll, A.H.; Yuan, X.; Pueschel, C.M. (2004). "Phosphatized multicellular algae in the Neoproterozoic Doushantuo Formation, China, and the early evolution of florideophyte red algae". American Journal of Botany 91 (2): 214–227. doi:10.3732/ajb.91.2.214. http://www.amjbot.org/cgi/content/full/91/2/214. 
  20. ^ Waggoner, Ben (1994–2008). "Introduction to the Phaeophyta: Kelps and brown "Algae"". University of California Museum of Palaeontology (UCMP). http://www.ucmp.berkeley.edu/chromista/phaeophyta.html. Retrieved 2008-12-19. 
  21. ^ a b Thomas, D N (2002). Seaweeds. London: The Natural History Museum. ISBN 0565091751. 
  22. ^ Waggoner, Ben (1994–2008). "Introduction to the Rhodophyta, The red "algae"". University of California Museum of Palaeontology (UCMP). http://www.ucmp.berkeley.edu/protista/rhodophyta.html. Retrieved 2008-12-19. 
  23. ^ Introduction to the Green Algae
  24. ^ Brodo, Irwin M; Sharnoff, Sylvia Duran; Sharnoff, Stephen; Laurie-Bourque, Susan (2001). Lichens of North America. New Haven: Yale University Press. p. 8. ISBN 0300082495, 9780300082494. 
  25. ^ Pearson, Lorentz C (1995). The Diversity and Evolution of Plants. CRC Press. p. 221. ISBN 0849324831, 9780849324833. 
  26. ^ Brodo et al. (2001), page 6: "A species of lichen collected anywhere in its range has the same lichen-forming fungus and, generally, the same photobiont. (A particular photobiont, on the other hand, may associate with scores of different lichen fungi)."
  27. ^ Brodo et al. (2001), page 8.
  28. ^ Taylor, Dennis L (1983). "The coral-algal symbiosis". in Goff, Lynda J. Algal Symbiosis: A Continuum of Interaction Strategies. CUP Archive. pp. 19–20. ISBN 0521255414, 9780521255417. 
  29. ^ http://uwsp.edu/cnr/UWEXlakes/laketides/vol26-4/vol26-4.pdf
  30. ^ Lobban, C S and Harrison, P J (1997) Seaweed Ecology and Physiology. Cambridge University Press. ISBN 0-521-40897-00
  31. ^ Algae II
  32. ^ "Algae Herbarium". National Museum of Natural History, Department of Botany. 2008. http://botany.si.edu/projects/algae/herbarium.htm. Retrieved 2008-12-19. 
  33. ^ John (2002), page 1.
  34. ^ Huisman (2000), page 25.
  35. ^ Stegenga (1997).
  36. ^ Abbott and Hollenberg (1976), page 2.
  37. ^ Hardy and Guiry (2006).
  38. ^ Round (1981), Chapter 8, Dispersal, continuity and phytogeography.
  39. ^ Round (1981), page 360.
  40. ^ Round (1981), page 362.
  41. ^ Round (1981), Page 357.
  42. ^ Round (1981), page 371.
  43. ^ Round (1981), page 366.
  44. ^ Round (1981), page 176.
  45. ^ Lewis, J G; Stanley, N F; Guist, G G (1988). "9 Commercial production of algal hydrocolloides". in Lembi, C.A.; Waaland, J.R.. Algae and Human Affairs. Cambridge University Press. ISBN 9780521321150. 
  46. ^ "Macrocystis C. Agardh 1820: 46". AlgaeBase. http://www.algaebase.org/generadetail.lasso?genus_id=35715&-session=abv3:51909EC307dcf25DFApmi3530315. Retrieved 2008-12-28. 
  47. ^ "Secondary Products of Brown Algae". Algae Research. Smithsonian National Museum of Natural History. http://botany.si.edu/projects/algae/economicuses/brownalgae.htm. Retrieved 2008-12-29. 
  48. ^ Read, Clare Sewell (1849). "On the Farming of South Wales: Prize Report". Journal of the Royal Agricultural Society of England (John Murray) 10: 142–143.  Downloadable Google Books.
  49. ^ McHugh, Dennis J. (2003). "9, Other Uses of Seaweeds". A Guide to the Seaweed Industry: FAO Fisheries Technical Paper 441. Rome: Fisheries and Aquaculture Department, Food and Agriculture Organization (FAO) of the United Nations. ISBN 9251049580, ISBN 0499345. http://www.fao.org/DOCREP/006/Y4765E/y4765e0c.htm#TopOfPage. 
  50. ^ Simoons, Frederick J (1991). "6, Seaweeds and Other Algae". Food in China: A Cultural and Historical Inquiry. CRC Press. pp. 179–190. ISBN 084938804X, 9780849388040. ISBN 0936923296. 
  51. ^ Morton, Steve L. "Modern Uses of Cultivated Algae". Ethnobotanical Leaflets. Southern Illinois University Carbondale. http://www.siu.edu/~ebl/leaflets/algae.htm. Retrieved 2008-12-26. 
  52. ^ Mondragon, J; Mondragon, J (2003). Seaweeds of the Pacific Coast. Monterey, California: Sea Challengers Publications. ISBN 0930118294. 
  53. ^ "Durvillaea antarctica (Chamisso) Hariot". AlgaeBase. http://www.algaebase.org/speciesdetail.lasso?species_id=11752&sk=0&from=results&-session=abv3:51909EC30802716127sVj3EDC9C7. 
  54. ^ Bigogno, C; I Khozin-Goldberg; S Boussiba; A Vonshak; Z Cohen (2002). "Lipid and fatty acid composition of the green oleaginous alga Parietochloris incisa, the richest plant source of arachidonic acid.". Phytochemistry 60 (5): 497–503. doi:10.1016/S0031-9422(02)00100-0. PMID 12052516. 
  55. ^ Allison Aubrey (Morning Edition, November 1, 2007). "Getting Brain Food Straight from the Source". National Public Radio. http://www.npr.org/templates/story/story.php?storyId=15823852. 
  56. ^ Arad, Shoshana; Spharim, Ishai (1998). "Production of Valuable Products from Microalgae: An Emerging Agroindustry". in Altman, Arie. Agricultural Biotechnology. Books in Soils, Plants, and the Environment. 61. CRC Press. p. 638. ISBN 0824794397, 9780824794392. 

Bibliography

General

  • Chapman, V.J. (1950). Seaweeds and their Uses. London: Methuen & Co. Ltd. ISBN 0412157403. 
  • Lembi, C.A.; Waaland, J.R. (1988). Algae and Human Affairs. Cambridge: Cambridge University Press. ISBN 0521321158. 
  • Round, F E (1981). The Ecology of Algae. London: Cambridge University Press. ISBN 0521225833. 
  • Mumford, T F; Miura, A (1988). "Porphyra as food: cultivation and economic". in Lembi, C A; Waaland, J R. Algae and Human Affairs. Cambridge University Press. pp. 87–117. ISBN 0521321158. .

Regional

Britain and Ireland
  • Brodie, Juliet; Burrows, Elsie M; Chamberlain, Yvonne M.; Christensen, Tyge; Dixon, Peter Stanley; Fletcher, R.L.; Hommersand, Max H; Irvine, Linda M et al. (1977–2003). Seaweeds of the British Isles: A Collaborative Project of the British Phycological Society and the British Museum (Natural History). London, Andover: British Museum (Natural History), HMSO, Intercept. ISBN 0565007815, 9780565007812, 0113100450, 9780113100453, 0113100167, 9780113100163, 1898298874, 9781898298878, 056500980X, 9780565009809, 0565009818, 0565009923, 0565008714. 
  • Cullinane, John P (1973). Phycology of the South Coast of Ireland. Cork: Cork University Press. 
  • Hardy, F G; Aspinall, R J (1988). An Atlas of the Seaweeds of Northumberland and Durham. The Hancock Museum, University Newcastle upon Tyne: Northumberland Biological Records Centre. ISBN 0950968056, 9780950968056. 
  • Hardy, F G; Guiry, Michael D; Arnold, Henry R (2006). A Check-list and Atlas of the Seaweeds of Britain and Ireland (Revised ed.). London: British Phycological Society. ISBN 390616635X 9783906166353. 
  • John, D M; Whitton, B A; Brook, J A (2002). The Freshwater Algal Flora of the British Isles. Cambridge, UK; New York: Cambridge University Press. ISBN 0521770513. 
  • Knight, Margery; Parke, Mary W (1931). Manx Algae: An Algal Survey of the South End of the Isle of Man. Liverpool Marine Biology Committee (LMBC) Memoirs on Typical British Marine Plants & Animals. XXX. Liverpool: University Press. 
  • Morton, Osborne (1994). Marine Algae of Northern Ireland. Belfast: Ulster Museum. ISBN 0900761288 9780900761287. 
  • Morton, Osborne (1 December 2003). "The Marine Macroalgae of County Donegal, Ireland". Bulletin of the Irish Biogeographical Society 27: 3–164. 
Australia
  • Huisman, J M (2000). Marine Plants of Australia. University of Western Australian (UWA) Press. ISBN 1876268336. 
New Zealand
  • Chapman, Valentine Jackson; Lindauer, VW; Aiken, M; Dromgoole, FI (1900, 1956, 1961, 1969, 1970). The Marine algae of New Zealand. London; Lehre, Germany: Linnaean Society of London; Cramer. 
Europe
  • Cabioc'h, Jacqueline; Floc'h, Jean-Yves; Le Toquin, Alain; Boudouresque, Charles-François; Meinesz, Alexandre; Verlaque, Marc (1992) (in French). Guide des algues des mers d'Europe: Manche/Atlantique-Méditerranée. Lausanne, Suisse: Delachaux et Niestlé. ISBN 260300848X, 9782603008485. 
  • Gayral, Paulette (1966) (in French). Les Algues de côtes françaises (manche et atlantique), notions fondamentales sur l'écologie, la biologie et la systématique des algues marines. Paris: Doin, Deren et Cie. 
  • Guiry, M.D.; Blunden, G. (1991). Seaweed Resources in Europe: Uses and Potential. John Wiley & Sons. ISBN 0471929476. 
  • Míguez Rodríguez, Luís (1998) (in Galician). Algas mariñas de Galicia: bioloxía, gastronomía, industria. Vigo: Edicións Xerais de Galicia. ISBN 84-8302-263-X. 
  • Otero, J. (2002) (in Galician). Guía das macroalgas de Galicia. A Coruña: Baía Edicións. ISBN 84-89803-22-6. 
  • Bárbara, I.; Cremades, J. (1993) (in Spanish). Guía de las algas del litoral gallego. A Coruña: Concello da Coruña - Casa das Ciencias. 
Arctic
  • Kjellman, Frans Reinhold (1883). The algae of the Arctic Sea: a survey of the species, together with an exposition of the general characters and the development of the flora. 20. Stockholm: Kungl. Svenska vetenskapsakademiens handlingar. pp. 1–350. 
Greenland
  • Lund, Søren Jensen (1959). The Marine Algae of East Greenland. Kövenhavn: C.A. Reitzel. ISBN 9584734. 
Faroe Islands
  • Børgesen, Frederik (1903, 1970 reprint). "Marine Algae". in Warming, Eugene. Botany of the Faröes Based Upon Danish Investigations. Part II. Det nordiske Forlag. pp. 339–532. .
Canary Islands
  • Børgesen, Frederik (1925, 1926, 1927, 1929, 1930, 1936). Marine Algae from the Canary Islands. København: Bianco Lunos. 
Morocco
  • Gayral, Paulette (1958) (in French). Algues de la côte atlantique marocaine. Casablanca: Rabat [Société des sciences naturelles et physiques du Maroc]. 
South Africa
  • Stegenga, H.; Bolton, J.J.; Anderson, R.J. (1997). Seaweeds of the South African West Coast. Bolus Herbarium, University of Cape Town. ISBN 079921793x. 
North America

External links



1911 encyclopedia

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Wiktionary

Up to date as of January 14, 2010
(Redirected to algae article)

Definition from Wiktionary, a free dictionary

Contents

English

Pronunciation

Noun

algae
  1. Plural form of alga.

Translations


Simple English

File:Dinobryon
Dinobryon, a colonial alga from the Chrysophyceae group

[[File:|200px|thumb|right|A seaweed (Laurencia) up close: the "branches" are multicellular and only about 1 mm thick. Much smaller algae are attached to the structure extending upwards in the lower right quarter]]

File:Phytoplankton SoAtlantic
Phytoplankton bloom in the south Atlantic off Argentina

Algae (one Alga, several Algae) is the name for certain plant-like living things that can make food from sunlight (photosynthesis). The study of algae is called phycology or algology.

Contents

Biology and taxonomy

Algae are a large and diverse group of simple, typically autotrophic organisms, ranging from unicellular to multicellular forms. The largest and most complex marine forms are called seaweeds. They are photosynthetic, like plants, and "simple" because they lack the many distinct organs found in land plants. For that reason they are not classified as plants.[1]

Though the prokaryotic Cyanobacteria (formerly referred to as blue-green algae) were included as "Algae" in older textbooks, this is outdated.[2] The term Algae is now used for eukaryotic organisms.[3] All true algae therefore have a nucleus enclosed within a membrane and chloroplasts bound in one or more membranes.[2][4] However, algae are definitely not a monophyletic group,[2] as they do not all descend from a common algal ancestor.[5] Modern taxonomists propose splitting them up into monophyletic groups, but there is at present no consensus as to the details.[5][6]

Algae lack the various structures that characterize land plants, such as leaves, roots, and other organs that are found in plants. Nearly all algae have photosynthetic machinery ultimately derived from the cyanobacteria, and so produce oxygen as a by-product of photosynthesis, unlike other photosynthetic bacteria such as purple and green bacteria. Some unicellular species rely entirely on external energy sources and have limited or no photosynthetic apparatus.

Fossilized filamentous algae from the Vindhya basin have been dated back to 1.6 to 1.7 billion years ago.[7]

Life style

Ecology

Algae are usually found in damp places or water, and are common on land as well as water. However, terrestrial algae are usually rather inconspicuous and are far more common in moist, tropical regions than dry ones. Algae lack vascular tissues and other adaptations to live on land, but they can endure dryness and other conditions in symbiosis with a fungus as lichen.

The various sorts of algae play significant roles in aquatic ecology. Microscopic forms that live suspended in the water column are called phytoplankton. They provide the food base for most marine food chains. Kelp grows mostly in shallow marine waters. Some are used as human food or harvested for agar or fertilizer. Kelp can grow in large stands called kelp forests. These forests prevent some of the damage from waves. Many different species live in them, including sea urchins, sea otters, and abalone.

Some algae may harm other species. Some algae may reproduce a lot, and make an algal bloom. These algae may produce protective toxins which can kill fish in the water. Dinoflagellates secrete a compound that turns the flesh of fish into slime. The algae then consume this nutritious liquid.

Symbiosis

File:Pteraeolidia ianthina Nick
Nudibranch Pteraeolidia ianthina contains algae which photosynthesize, and provide food for the mollusc

Algae have evolved a number of symbiotic partnerships with other organisms. The most famous is the plant-like lichen, which are each formed by a fungus with an alga. It is a highly successful life-form, and twenty thousand 'species' are known. In all cases the lichen are quite different in appearance and life-style from either constituent; it is possibly the most complete symbiosis known. Both constituents gain from their access to niches with low nutrient value, which is where lichen are found.

Less well known are the algal relationships with animals. Reef-building corals are basically social Cnidarian polyps. Corals are dependent on light, because the algae are important partners, and they require light. Corals have evolved structures, often tree-like, which offer the algae maximum access to light. The coral weakens the algal cell walls, and digests about 80% of the food synthesised by the algae. The corals' waste-products provide nutrients for the algae so, as with lichen, both partners gain from the association. The algae are golden-brown flagellate algae, often of the genus Symbiodinium. A curious feature of the partnership is that the coral may eject the algae in hard times, and regain them later. The ejection of the algal partner is called bleaching, because the coral loses its colour.[8]p200

Other types of Cnideria, such as sea anemones and jellyfish, also contain algae. Jellyfish with algae behave so that their partners get the best light during the day, and descend to depths at night, where the water is rich in nitrates and brown with decay. Sea slugs and clams are also well-known for harbouring algae. Both groups are molluscs. The sea slugs graze on coral, and are the same colour as the coral they graze. They are able to separate the algae from the polyp tissues they digest. The algal cells are moved to its tentacles, where they continue to live. The otherwise defenceless slug gains both camouflage and nutrition.[8]p204 The giant clam keeps algae in its mantle, which is revealed when the clam is open. The coloured mantle has places where the skin is transparent, and acts like a lens to concentrate light on the algae beneath. When the algae get too numerous, the clam digests them.[8]p203

Various other marine invertebrate groups have members which symbiont with algae. Flatworms (Platyhelminths) and Polychaete worms (Annelids) are two such groups.

References

  1. The American Heritage Dictionary of the English Language: 4th ed, 2000.
  2. 2.0 2.1 2.2 Nabors, Murray W. (2004). Introduction to Botany. San Francisco, CA: Pearson Education, Inc. 
  3. Allaby, M ed. (1992). "Algae". The Concise Dictionary of Botany. Oxford: Oxford University Press. 
  4. Round (1981).
  5. 5.0 5.1 Patrick J. Keeling (2004). "Diversity and evolutionary history of plastids and their hosts". American Journal of Botany 91: 1481–1493. doi:10.3732/ajb.91.10.1481. http://www.amjbot.org/cgi/content/full/91/10/1481. 
  6. Laura Wegener Parfrey, Erika Barbero, Elyse Lasser, Micah Dunthorn, Debashish Bhattacharya, David J Patterson, and Laura A Katz (December 2006). "Evaluating Support for the Current Classification of Eukaryotic Diversity". PLoS Genet. 2 (12): e220. doi:10.1371/journal.pgen.0020220. PMID 17194223. PMC 1713255. http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=1713255. 
  7. Bengtson S, Belivanova V, Rasmussen B, Whitehouse M. 2009. The controversial "Cambrian" fossils of the Vindhyan are real but more than a billion years older. PNAS 106:7729-34.
  8. 8.0 8.1 8.2 Attenborough, David. 1995. The private life of plants. BBC, London. Chapter 5 Living together.


Citable sentences

Up to date as of December 29, 2010

Here are sentences from other pages on Algae, which are similar to those in the above article.








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