Algae
Algae An informal term for a diverse group of photosynthetic eukaryotes Temporal range:
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A variety of algae growing on the sea bed in shallow waters | |
A variety of microscopic unicellular and colonial freshwater algae | |
Scientific classification | |
Domain: | Eukaryota |
Groups included | |
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Typically excluded | |
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Algae (
Algae constitute a polyphyletic group
Algae lack the various structures that characterize
Because of the wide range of types of algae, they have increasing different industrial and traditional applications in human society. Traditional seaweed farming practices have existed for thousands of years and have strong traditions in East Asia food cultures. More modern algaculture applications extend the food traditions for other applications include cattle feed, using algae for bioremediation or pollution control, transforming sunlight into algae fuels or other chemicals used in industrial processes, and in medical and scientific applications. A 2020 review found that these applications of algae could play an important role in carbon sequestration in order to mitigate climate change while providing lucrative value-added products for global economies.[12]
Etymology and study
The singular alga is the Latin word for 'seaweed' and retains that meaning in English.[13] The etymology is obscure. Although some speculate that it is related to Latin algēre, 'be cold',[14] no reason is known to associate seaweed with temperature. A more likely source is alliga, 'binding, entwining'.[15]
The Ancient Greek word for 'seaweed' was φῦκος (phŷkos), 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, or blue.[16]
The study of algae is most commonly called phycology (from Greek phykos 'seaweed'); the term algology is falling out of use.[17]
Classifications
One definition of algae is that they "have chlorophyll as their primary photosynthetic pigment and lack a sterile covering of cells around their reproductive cells".[18] On the other hand, the colorless Prototheca under Chlorophyta are all devoid of any chlorophyll. Although cyanobacteria are often referred to as "blue-green algae", most authorities exclude all prokaryotes, including cyanobacteria, from the definition of algae.[4][19]
The algae contain chloroplasts that are similar in structure to cyanobacteria. Chloroplasts contain circular DNA like that in cyanobacteria and are interpreted as representing reduced endosymbiotic cyanobacteria. However, the exact origin of the chloroplasts is different among separate lineages of algae, reflecting their acquisition during 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.[20]
not nucleocytoplasmic genealogy:
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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 without phycobilins. Land plants are pigmented similarly to green algae and probably developed from them, thus the Chlorophyta is a sister taxon to the plants; sometimes the Chlorophyta, the Charophyta, and land plants are grouped together as the Viridiplantae. |
Excavata and Rhizaria | Green algae |
These groups have green chloroplasts containing chlorophylls a and b.[22] 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 fresh water and have chloroplasts with only three membranes. The endosymbiotic green algae may have been acquired through myzocytosis rather than phagocytosis.[23] (Another group with green algae endosymbionts is the dinoflagellate genus Lepidodinium, which has replaced its original endosymbiont of red algal origin with one of green algal origin. A nucleomorph is present, and the host genome still have several red algal genes acquired through endosymbiotic gene transfer. Also the euglenid and chlorarachniophyte genome contain genes of apparent red algal ancestry)[24][25][26] | |
Halvaria and Hacrobia |
|
Red algae |
These groups have chloroplasts containing chlorophylls a and c, and phycobilins. The shape can vary; they may be of discoid, plate-like, reticulate, cup-shaped, spiral, or ribbon shaped. They have one or more pyrenoids to preserve protein and starch. The latter chlorophyll type is not known from any prokaryotes or primary chloroplasts, but genetic similarities with red algae suggest a relationship there.[27] In the first three of these groups ( Haptophyta, and cryptomonads are in fact more closely related to each other than to other groups.[28][29]
The typical dinoflagellate chloroplast has three membranes, but considerable diversity exists in chloroplasts within the group, and a number of endosymbiotic events apparently occurred.[5] The Apicomplexa, a group of closely related parasites, also have plastids called apicoplasts, which are not photosynthetic, but appear to have a common origin with dinoflagellate chloroplasts.[5] |
Linnaeus, in Species Plantarum (1753),[30] the starting point for modern botanical nomenclature, recognized 14 genera of algae, of which only four are currently considered among algae.[31] In Systema Naturae, Linnaeus described the genera Volvox and Corallina, and a species of Acetabularia (as Madrepora), among the animals.
In 1768, Samuel Gottlieb Gmelin (1744–1774) published the Historia Fucorum, the first work dedicated to marine algae and the first book on marine biology to use the then new binomial nomenclature of Linnaeus. It included elaborate illustrations of seaweed and marine algae on folded leaves.[32][33]
At this time, microscopic algae were discovered and reported by a different group of workers (e.g.,
Although used as a taxonomic category in some pre-Darwinian classifications, e.g., Linnaeus (1753), de Jussieu (1789), Horaninow (1843), Agassiz (1859), Wilson & Cassin (1864), in further classifications, the "algae" are seen as an artificial, polyphyletic group.[citation needed]
Throughout the 20th century, most classifications treated the following groups as divisions or classes of algae:
With the abandonment of plant-animal dichotomous classification, most groups of algae (sometimes all) were included in
Some parasitic algae (e.g., the green algae
Evolution
Algae are
Recent
Relationship to land plants
Fossils of isolated
Morphology
A range of algal
Most of the simpler algae are unicellular
- Colonial: small, regular groups of motile cells
- Capsoid: individual non-motile cells embedded in mucilage
- Coccoid: individual non-motile cells with cell walls
- Palmelloid: nonmotile cells embedded in mucilage
- Filamentous: a string of connected nonmotile cells, 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,[50]—some of which may reach 50 m in length (kelps)[51]—the red algae,[52] and the green algae.[53] The most complex forms are found among the charophyte algae (see Charales and Charophyta), in a lineage that eventually led to the higher land plants. The innovation that defines these nonalgal plants is the presence of female reproductive organs with protective cell layers that protect the zygote and developing embryo. Hence, the land plants are referred to as the Embryophytes.
Turfs
The term algal turf is commonly used but poorly defined. Algal turfs are thick, carpet-like beds of seaweed that retain sediment and compete with foundation species like corals and kelps, and they are usually less than 15 cm tall. Such a turf may consist of one or more species, and will generally cover an area in the order of a square metre or more. Some common characteristics are listed:[54]
- Algae that form aggregations that have been described as turfs include diatoms, cyanobacteria, chlorophytes, phaeophytes and rhodophytes. Turfs are often composed of numerous species at a wide range of spatial scales, but monospecific turfs are frequently reported.[54]
- Turfs can be morphologically highly variable over geographic scales and even within species on local scales and can be difficult to identify in terms of the constituent species.[54]
- Turfs have been defined as short algae, but this has been used to describe height ranges from less than 0.5 cm to more than 10 cm. In some regions, the descriptions approached heights which might be described as canopies (20 to 30 cm).[54]
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Physiology
Many algae, particularly species of the
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:
Lichens
Coral reefs
Sea sponges
Endosymbiontic green algae live close to the surface of some sponges, for example, breadcrumb sponges (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.[62]
Life cycle
Numbers
The Algal Collection of the US National Herbarium (located in the National Museum of Natural History) consists of approximately 320,500 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).[67] Estimates vary widely. For example, according to one standard textbook,[68] in the British Isles the UK Biodiversity Steering Group Report estimated there to be 20,000 algal species in the UK. Another checklist reports only about 5,000 species. Regarding the difference of about 15,000 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:
- 5,000–5,500 species of red algae worldwide
- "some 1,300 in Australian Seas"[69]
- 400 seaweed species for the western coastline of South Africa,[70] and 212 species from the coast of KwaZulu-Natal.[71] Some of these are duplicates, as the range extends across both coasts, and the total recorded is probably about 500 species. Most of these are listed in List of seaweeds of South Africa. These exclude phytoplankton and crustose corallines.
- 669 marine species from California (US)[72]
- 642 in the check-list of Britain and Ireland[73]
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 microscopic algae, such as phytoplankton.
The most recent estimate suggests 72,500 algal species worldwide.[74]
Distribution
The distribution of algal species has been fairly well studied since the founding of phytogeography in the mid-19th century.[75] Algae spread mainly by the dispersal of spores analogously to the dispersal of cryptogamic plants by spores. Spores can be found in a variety of environments: fresh and marine waters, air, soil, and in or on other organisms.[75] Whether a spore is to grow into an adult organism depends on the species and the environmental conditions where the spore lands.
The spores of freshwater algae are dispersed mainly by running water and wind, as well as by living carriers.[75] However, not all bodies of water can carry all species of algae, as the chemical composition of certain water bodies limits the algae that can survive within them.[75] Marine spores are often spread by ocean currents. Ocean water presents many vastly different habitats based on temperature and nutrient availability, resulting in phytogeographic zones, regions, and provinces.[76]
To some degree, the distribution of algae is subject to floristic discontinuities caused by geographical features, such as
Mapping is possible for select species only: "there are many valid examples of confined distribution patterns."[77] For example, Clathromorphum is an arctic genus and is not mapped far south of there.[78] However, scientists regard the overall data as insufficient due to the "difficulties of undertaking such studies."[79]
Ecology
Algae are prominent in bodies of water, common in terrestrial environments, and are found in unusual environments, such as on
The various sorts of algae play significant roles in aquatic ecology. Microscopic forms that live suspended in the water column (
Algae can be used as indicator organisms to monitor pollution in various aquatic systems.[83] In many cases, algal metabolism is sensitive to various pollutants. Due to this, the species composition of algal populations may shift in the presence of chemical pollutants.[83] To detect these changes, algae can be sampled from the environment and maintained in laboratories with relative ease.[83]
On the basis of their habitat, algae can be categorized as:
Cultural associations
In classical Chinese, the word 藻 is used both for "algae" and (in the modest tradition of the imperial scholars) for "literary talent". The third island in Kunming Lake beside the Summer Palace in Beijing is known as the Zaojian Tang Dao (藻鑒堂島), which thus simultaneously means "Island of the Algae-Viewing Hall" and "Island of the Hall for Reflecting on Literary Talent".
Cultivation
Algaculture is a form of aquaculture involving the farming of species of algae.[87]
The majority of algae that are intentionally cultivated fall into the category of
Commercial and industrial algae cultivation has numerous uses, including production of
Global production of farmed aquatic plants, overwhelmingly dominated by seaweeds, grew in output volume from 13.5 million tonnes in 1995 to just over 30 million tonnes in 2016.[93] Cultured microalgae already contribute to a wide range of sectors in the emerging bioeconomy.[94] Research suggests there are large potentials and benefits of algaculture for the development of a future healthy and sustainable food system.[95][92]Seaweed farming
Seaweed farming or kelp farming is the practice of cultivating and harvesting seaweed. In its simplest form farmers gather from natural beds, while at the other extreme farmers fully control the crop's life cycle.
The seven most cultivated
The largest seaweed-producing countries as of 2022 are China (58.62%) and Indonesia (28.6%); followed by South Korea (5.09%) and the Philippines (4.19%). Other notable producers include North Korea (1.6%), Japan (1.15%), Malaysia (0.53%), Zanzibar (Tanzania, 0.5%), and Chile (0.3%).[98][99] Seaweed farming has frequently been developed to improve economic conditions and to reduce fishing pressure.[100]
The Food and Agriculture Organization (FAO), reported that world production in 2019 was over 35 million tonnes. North America produced some 23,000 tonnes of wet seaweed. Alaska, Maine, France, and Norway each more than doubled their seaweed production since 2018. As of 2019, seaweed represented 30% of marine aquaculture.[101]
Seaweed farming is aBioreactors
Uses
Agar
Agar, a gelatinous substance derived from red algae, has a number of commercial uses.[106] It is a good medium on which to grow bacteria and fungi, as most microorganisms cannot digest agar.
Alginates
Between 100,000 and 170,000 wet tons of Macrocystis are harvested annually in New Mexico for alginate extraction and abalone feed.[107][108]
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,[109][110] 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 is estimated to occur by 2025.[111]
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:[112]
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.[113] 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
Naturally growing seaweeds are an important source of food, especially in Asia, leading some to label them as
Algae are national foods of many nations: China consumes more than 70 species, including
Two popular forms of algae are used in cuisine:
- vitamin B2 and Omega-3 fatty acids.
Furthermore, it contains all nine of the essential amino acids the body does not produce on its own[122]
- Spirulina: Known otherwise as a cyanobacterium (a prokaryote or a "blue-green alga")
The oils from some algae have high levels of
Pollution control
- Sewage can be treated with algae,[125] reducing the use of large amounts of toxic chemicals that would otherwise be needed.
- Algae can be used to capture fertilizers in runoff from farms. When subsequently harvested, the enriched algae can be used as fertilizer.
- Aquaria and ponds can be filtered using algae, which absorb nutrients from the water in a device called an algae scrubber, also known as an algae turf scrubber.[126][127]
Polymers
Various polymers can be created from algae, which can be especially useful in the creation of bioplastics. These include hybrid plastics, cellulose-based plastics, poly-lactic acid, and bio-polyethylene.[129] Several companies have begun to produce algae polymers commercially, including for use in flip-flops[130] and in surf boards.[131]
Bioremediation
The alga Stichococcus bacillaris has been seen to colonize silicone resins used at archaeological sites; biodegrading the synthetic substance.[132]
Pigments
The natural pigments (carotenoids and chlorophylls) produced by algae can be used as alternatives to chemical dyes and coloring agents.[133] The presence of some individual algal pigments, together with specific pigment concentration ratios, are taxon-specific: analysis of their concentrations with various analytical methods, particularly high-performance liquid chromatography, can therefore offer deep insight into the taxonomic composition and relative abundance of natural algae populations in sea water samples.[134][135]
Stabilizing substances
Carrageenan, from the red alga Chondrus crispus, is used as a stabilizer in milk products.
Additional images
-
Algae bladder
See also
- AlgaeBase
- AlgaePARC
- Eutrophication
- Iron fertilization
- Marimo algae
- Microbiofuels
- Microphyte
- Photobioreactor
- Phycotechnology
- Plant
- Toxoid – anatoxin
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Bibliography
General
- Chapman, V.J. (1950). Seaweeds and their Uses. London: Methuen. ISBN 978-0-412-15740-0.
- Fritsch, F. E. (1945) [1935]. The Structure and Reproduction of the Algae. Vol. I & II. Cambridge University Press.
- van den Hoek, C.; Mann, D. G.; Jahns, H. M. (1995). Algae: An Introduction to Phycology. Cambridge University Press.
- Kassinger, Ruth (2020). Slime: How Algae Created Us, Plague Us, and Just Might Save Us. Mariner.
- Lembi, C. A.; Waaland, J.R. (1988). Algae and Human Affairs. Cambridge University Press. ISBN 978-0-521-32115-0.
- Mumford, T. F.; Miura, A. (1988). "Porphyra as food: cultivation and economic". In Lembi, C. A.; Waaland, J. R. (eds.). Algae and Human Affairs. Cambridge University Press. pp. 87–117. ISBN 978-0-521-32115-0..
- Round, F. E. (1981). The Ecology of Algae. London: Cambridge University Press. ISBN 978-0-521-22583-0.
- Smith, G. M. (1938). Cryptogamic Botany. Vol. I. New York: McGraw-Hill.
- Ask, E.I (1990). Cottonii and Spinosum Cultivation Handbook. FMC BioPolymer Corporation.Philippines.
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.; Maggs, Christine A. (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 of Natural History, HMSO / Intercept. ISBN 978-0-565-00781-2.
- 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 978-0-9509680-5-6.
- Hardy, F. G.; ISBN 978-3-906166-35-3.
- John, D. M.; Whitton, B. A.; Brook, J. A. (2002). The Freshwater Algal Flora of the British Isles. Cambridge / New York: Cambridge University Press. ISBN 978-0-521-77051-4.
- Knight, Margery; Parke, Mary W. (1931). Manx Algae: An Algal Survey of the South End of the Isle of Man. Liverpool Marine Biology Committee Memoirs on Typical British Marine Plants & Animals. Vol. XXX. Liverpool: University Press.
- Morton, Osborne (1994). Marine Algae of Northern Ireland. Belfast: Ulster Museum. ISBN 978-0-900761-28-7.
- 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 Australia Press. ISBN 978-1-876268-33-6.
New Zealand
- Chapman, Valentine Jackson; Lindauer, VW; Aiken, M.; Dromgoole, F. I. (1970) [1900, 1956, 1961, 1969]. The Marine algae of New Zealand. London / Lehre, Germany: Linnean Society of London / Cramer.
Europe
- Cabioc'h, Jacqueline; Floc'h, Jean-Yves; Le Toquin, Alain; Boudouresque, Charles-François; Meinesz, Alexandre; Verlaque, Marc (1992). Guide des algues des mers d'Europe: Manche/Atlantique-Méditerranée (in French). Lausanne, Suisse: Delachaux et Niestlé. ISBN 978-2-603-00848-5.
- Gayral, Paulette (1966). Les Algues de côtes françaises (manche et atlantique), notions fondamentales sur l'écologie, la biologie et la systématique des algues marines (in French). Paris: Doin, Deren et Cie.
- ISBN 978-0-471-92947-5.
- Míguez Rodríguez, Luís (1998). Algas mariñas de Galicia: Bioloxía, gastronomía, industria (in Galician). Vigo: Edicións Xerais de Galicia. ISBN 978-84-8302-263-4.
- Otero, J. (2002). Guía das macroalgas de Galicia (in Galician). A Coruña: Baía Edicións. ISBN 978-84-89803-22-0.
- Bárbara, I.; Cremades, J. (1993). Guía de las algas del litoral gallego (in Spanish). 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. Vol. 20. Stockholm: Kungl. Svenska vetenskapsakademiens handlingar. pp. 1–350.
Greenland
- Lund, Søren Jensen (1959). The Marine Algae of East Greenland. Kövenhavn: C.A. Reitzel. 9584734.
Faroe Islands
- Børgesen, Frederik (1970) [1903]. "Marine Algae". In Warming, Eugene (ed.). Botany of the Faröes Based Upon Danish Investigations, Part II. Copenhagen: Det nordiske Forlag. pp. 339–532..
Canary Islands
- Børgesen, Frederik (1936) [1925, 1926, 1927, 1929, 1930]. Marine Algae from the Canary Islands. Copenhagen: Bianco Lunos.
Morocco
- Gayral, Paulette (1958). Algues de la côte atlantique marocaine (in French). 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 978-0-7992-1793-3.
North America
- Abbott, I. A.; Hollenberg, G. J. (1976). Marine Algae of California. California: Stanford University Press. ISBN 978-0-8047-0867-8.
- Greeson, Phillip E. (1982). An annotated key to the identification of commonly occurring and dominant genera of Algae observed in the Phytoplankton of the United States. Washington DC: US Department of the Interior, Geological Survey. Retrieved 19 December 2008.
- Taylor, William Randolph (1969) [1937, 1957, 1962]. Marine Algae of the Northeastern Coast of North America. Ann Arbor: University of Michigan Press. ISBN 978-0-472-04904-2.
- Wehr, J. D.; Sheath, R. G. (2003). Freshwater Algae of North America: Ecology and Classification. Academic Press. ISBN 978-0-12-741550-5.
External links
- Guiry, Michael; Guiry, Wendy. "AlgaeBase". – a database of all algal names including images, nomenclature, taxonomy, distribution, bibliography, uses, extracts
- "Algae – Cell Centered Database". CCDb.UCSD.edu. San Diego: University of California.
- Anderson, Don; Keafer, Bruce; Kleindinst, Judy; Shaughnessy, Katie; Joyce, Katherine; Fino, Danielle; Shepherd, Adam (2007). "Harmful Algae". US National Office for Harmful Algal Blooms. Archived from the original on 5 December 2008. Retrieved 19 December 2008.
- "About Algae". NMH.ac.uk. Natural History Museum, United Kingdom.