Algae (; singular alga ) is an informal term for a large, diverse group of photosynthetic organisms that are not necessarily closely related, and is thus polyphyletic. Included organisms range from unicellular microalgae genera, such as Chlorella and the diatoms, to multicellular forms, such as the giant kelp, a large brown alga which may grow up to 50 m in length. Most are aquatic and autotrophic and lack many of the distinct cell and tissue types, such as stomata, xylem, and phloem, which are found in land plants. The largest and most complex marine algae are called seaweeds, while the most complex freshwater forms are the Charophyta, a division of green algae which includes, for example, Spirogyra and the stoneworts. No definition of algae is generally accepted. One definition is that algae "have chlorophyll as their primary photosynthetic pigment and lack a sterile covering of cells around their reproductive cells". Some authors exclude all prokaryotes and thus do not consider cyanobacteria (blue-green algae) as algae. Algae constitute a polyphyletic group since they do not include a common ancestor, and although their plastids seem to have a single origin, from cyanobacteria, they were acquired in different ways. Green algae are examples of algae that have primary chloroplasts derived from endosymbiotic cyanobacteria. Diatoms and brown algae are examples of algae with secondary chloroplasts derived from an endosymbiotic red alga. Algae exhibit a wide range of reproductive strategies, from simple asexual cell division to complex forms of sexual reproduction.Smithsonian National Museum of Natural History; Department of Botany. Algae lack the various structures that characterize land plants, such as the phyllids (leaf-like structures) of bryophytes, rhizoids in nonvascular plants, and the roots, leaves, and other organs found in tracheophytes ( vascular plants). Most are phototrophic, although some are mixotrophic, deriving energy both from photosynthesis and uptake of organic carbon either by osmotrophy, myzotrophy, or phagotrophy. Some unicellular species of green algae, many golden algae, euglenids, dinoflagellates, and other algae have become heterotrophs (also called colorless or apochlorotic algae), sometimes parasitic, relying entirely on external energy sources and have limited or no photosynthetic apparatus.Pringsheim, E.G. 1963. Farblose Algen. Ein beitrag zur Evolutionsforschung. Gustav Fischer Verlag, Stuttgart. 471 pp., . Some other heterotrophic organisms, such as the apicomplexans, are also derived from cells whose ancestors possessed plastids, but are not traditionally considered as algae. Algae have photosynthetic machinery ultimately derived from cyanobacteria that 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.
Etymology and studyThe singular alga is the Latin word for "seaweed" and retains that meaning in English. The etymology is obscure. Although some speculate that it is related to Latin algēre, "be cold", no reason is known to associate seaweed with temperature. A more likely source is alliga, "binding, entwining". The Ancient Greek word for seaweed was φῦκος (phŷcos), 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. Accordingly, the modern study of marine and freshwater algae is called either phycology or algology, depending on whether the Greek or Latin root is used. The name Fucus appears in a number of taxa.
Classificationof the unicellular coccolithophore Gephyrocapsa oceanica]] Most algae contain chloroplasts that are similar in structure to cyanobacteria. Chloroplasts contain circular DNA like that in cyanobacteria and presumably represent 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. Phylogeny based on plastid not nucleocytoplasmic genealogy: |2= |2= Chlorarachniophytes }} }} }} }} }} Historia Fucorum, dated 1768]] Linnaeus, in Species Plantarum (1753),Linnæus, C. (1753). Species Plantarum, vol. 2, p. 1131, link . the starting point for modern botanical nomenclature, recognized 14 genera of algae, of which only four are currently considered among algae.Sharma, O. P. (1986). Textbook of Algae. McGraw Hill. p. 22, . 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.Gmelin S G (1768) Historia Fucorum Ex typographia Academiae scientiarum, St. Petersburg.Silva PC, Basson PW and Moe RL (1996) Catalogue of the Benthic Marine Algae of the Indian Ocean page 2, University of California Press. . W.H.Harvey (1811—1866) and Lamouroux (1813)Medlin, L. K., W. H. C. F. Kooistra, D. Potter, G. W. Saunders, and R. A. Anderson (1997). Phylogenetic relationships of the "golden algae" (haptophytes, heterokont chromophytes) and their plastids. Plant Systematics and Evolution, p. 188. were the first to divide macroscopic 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 (Rhodospermae), brown algae (Melanospermae), green algae (Chlorospermae), and Diatomaceae.Harvey, D. (1836). Flora hibernica. At this time, microscopic algae were discovered and reported by a different group of workers (e.g., O. F. Müller and Ehrenberg) studying the Infusoria (microscopic organisms). Unlike macroalgae, which were clearly viewed as plants, microalgae were frequently considered animals because they are often motile.Medlin et al. (1997), p. 188. Even the nonmotile (coccoid) microalgae were sometimes merely seen as stages of the lifecycle of plants, macroalgae, or animals.Braun, A. Algarum unicellularium genera nova et minus cognita, praemissis observationibus de algis unicellularibus in genere (New and less known genera of unicellular algae, preceded by observations respecting unicellular algae in general) . Lipsiae, Apud W. Engelmann, 1855. Translation at: Lankester, E. & Busk, G. (eds.). Quarterly Journal of Microscopical Science, 1857, vol. 5, (17), 13-16 ; (18), 90-96 ; (19), 143-149 .Siebold, C. Th. v. " Ueber einzellige Pflanzen und Thiere (On unicellular plants and animals) ". In: Siebold, C. Th. v. & Kölliker, A. (1849). Zeitschrift für wissenschaftliche Zoologie, Bd. 1, p. 270. Translation at: Lankester, E. & Busk, G. (eds.). Quarterly Journal of Microscopical Science, 1853, vol. 1, (2), 111-121 ; (3), 195-206 . 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. Throughout the 20th century, most classifications treated the following groups as divisions or classes of algae: cyanophytes, rhodophytes, chrysophytes, xanthophytes, bacillariophytes, phaeophytes, pyrrhophytes ( cryptophytes and dinophytes), euglenophytes, and chlorophytes. Later, many new groups were discovered (e.g., Bolidophyceae), and others were splintered from older groups: charophytes and glaucophytes (from chlorophytes), many heterokontophytes (e.g., synurophytes from chrysophytes, or eustigmatophytes from xanthophytes), haptophytes (from chrysophytes), and chlorarachniophytes (from xanthophytes). With the abandonment of plant-animal dichotomous classification, most groups of algae (sometimes all) were included in Protista, later also abandoned in favour of Eukaryota. However, as a legacy of the older plant life scheme, some groups that were also treated as protozoans in the past still have duplicated classifications (see ambiregnal protists). Some parasitic algae (e.g., the green algae Prototheca and Helicosporidium, parasites of metazoans, or Cephaleuros, parasites of plants) were originally classified as fungi, sporozoans, or protistans of incertae sedis, while others (e.g., the green algae Phyllosiphon and Rhodochytrium, parasites of plants, or the red algae Pterocladiophila and Gelidiocolax mammillatus, parasites of other red algae, or the dinoflagellates Oodinium, parasites of fish) had their relationship with algae conjectured early. In other cases, some groups were originally characterized as parasitic algae (e.g., Chlorochytrium), but later were seen as endophytic algae.Round (1981). pp. 398–400, . Some filamentous bacteria (e.g., Beggiatoa) were originally seen as algae. Furthermore, groups like the apicomplexans are also parasites derived from ancestors that possessed plastids, but are not included in any group traditionally seen as algae.
Relationship to land plantsThe first land plants probably evolved from shallow freshwater charophyte algae much like Chara almost 500 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.
Morphologyexhibit at the Monterey Bay Aquarium: A three-dimensional, multicellular thallus]] A range of algal morphologies is 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. Apical growth is constrained to subsets of these groups: the florideophyte reds, various browns, and the charophytes. The form of charophytes is quite different from those of reds and browns, because they have distinct nodes, separated by internode 'stems'; whorls of branches reminiscent of the horsetails occur at the nodes. Conceptacles are another polyphyletic trait; they appear in the coralline algae and the Hildenbrandiales, as well as the browns. Most of the simpler algae are unicellular flagellates or amoeboids, but colonial and nonmotile 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 lifecycle 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: nonmotile cells embedded in mucilage
- Filamentous: a string of nonmotile cells connected together, sometimes branching
- Parenchymatous: cells forming a thallus with partial differentiation of tissues
PhysiologyMany algae, particularly members of the Characeae, have served as model experimental organisms to understand the mechanisms of the water permeability of membranes, osmoregulation, turgor regulation, salt tolerance, cytoplasmic streaming, and the generation of action potentials. Phytohormones are found not only in higher plants, but in algae, too.
Symbiotic algaeSome 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:
LichensLichens 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." The fungi, or mycobionts, are mainly 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. One mycobiont associates with the same phycobiont species, rarely two, from the green algae, except that alternatively, the mycobiont may associate with a species of cyanobacteria (hence "photobiont" is the more accurate term). A photobiont may be associated with many different mycobionts or may live independently; accordingly, lichens are named and classified as fungal species.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, though, may associate with scores of different lichen fungi)." 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). The photobiont possibly triggers otherwise latent genes in the mycobiont.Brodo et al. (2001), page 8.
Coral reefsCoral reefs are accumulated from the calcareous exoskeletons of marine invertebrates of the order Scleractinia (stony corals). These animals metabolize sugar and oxygen to obtain energy for their cell-building processes, including secretion of the exoskeleton, with water and carbon dioxide as byproducts. Dinoflagellates (algal protists) are often endosymbionts in the cells of the coral-forming 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 by the host. Reef-building stony corals ( hermatypic corals) require endosymbiotic algae from the genus Symbiodinium to be in a healthy condition. The loss of Symbiodinium from the host is known as coral bleaching, a condition which leads to the deterioration of a reef.
Sea spongesGreen 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.
LifecycleRhodophyta, Chlorophyta, and Heterokontophyta, the three main algal divisions, have lifecycles which show considerable variation and complexity. In general, an asexual phase exists 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 permits efficient population increases, but less variation is possible. Commonly, in sexual reproduction of unicellular and colonial algae, two specialized, sexually compatible, haploid gametes make physical contact and fuse to form a zygote. To ensure a successful mating, the development and release of gametes is highly synchronized and regulated; pheromones may play a key role in these processes. Sexual reproduction allows for more variation and provides the benefit of efficient recombinational repair of DNA damages during meiosis, a key stage of the sexual cycle.Harris Bernstein, Carol Bernstein and Richard E. Michod (2011). Meiosis as an Evolutionary Adaptation for DNA Repair. Chapter 19 pages 357–382 in "DNA Repair" (Inna Kruman editor). InTech Open Publisher. However, sexual reproduction is more costly than asexual reproduction. Meiosis has been shown to occur in many different species of algae.
Numbersin Taiwan]] 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). Estimates vary widely. For example, according to one standard textbook,John (2002), page 1. 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"Huisman (2000), page 25.
- 400 seaweed species for the western coastline of South Africa,Stegenga (1997). and 212 species from the coast of KwaZulu-Natal. 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)Abbott and Hollenberg (1976), page 2.
- 642 in the check-list of Britain and IrelandHardy and Guiry (2006).
DistributionThe distribution of algal species has been fairly well studied since the founding of phytogeography in the mid-19th century. Algae spread mainly by the dispersal of spores analogously to the dispersal of Plantae by seeds and spores. This dispersal can be accomplished by air, water, or other organisms. Due to this, spores can be found in a variety of environments: fresh and marine waters, air, soil, and in or on other organisms. Whether a spore is to grow into an organism depends on the combination of 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. 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. 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.Round (1981), page 362. 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, hypothesizing a transport mechanism is usually possible, such as the hulls of ships. For example, Ulva reticulata and U. 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."Round (1981), Page 357. For example, Clathromorphum is an arctic genus and is not mapped far south of there.Round (1981), page 371. However, scientists regard the overall data as insufficient due to the "difficulties of undertaking such studies."Round (1981), page 366.
Ecology]] Algae are prominent in bodies of water, common in terrestrial environments, and are found in unusual environments, such as on snow and ice. Seaweeds grow mostly in shallow marine waters, under deep; however, some such as Navicula pennata have been recorded to a depth of .Round (1981), page 176. 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 can be used as indicator organisms to monitor pollution in various aquatic systems. 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. To detect these changes, algae can be sampled from the environment and maintained in laboratories with relative ease. On the basis of their habitat, algae can be categorized as: aquatic ( planktonic, benthic, marine, freshwater, lentic, lotic),Necchi Jr., O. (ed.) (2016). River Algae. Springer, . terrestrial, aerial (subareial),Johansen, J. R. 2012. Diatoms of aerial habitats. In: Smol, J. P. & Stoermer, E. F. (Eds.). The Diatoms: Applications for the Environmental and Earth Sciences, 2nd ed. Cambridge University Press. Cambridge, UK. pp. 465–472, link . lithophytic, halophytic (or euryhaline), psammon, thermophilic, cryophilic, epibiont ( epiphytic, epizoic), endosymbiont ( endophytic, endozoic), parasitic, calcifilic or lichenic (phycobiont).Sharma, O. P. (1986). pp. 2–6, link .
Cultural associationsIn classical Chinese, the word }} 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, which thus simultaneously means "Island of the Algae-Viewing Hall" and "Island of the Hall for Reflecting on Literary Talent".
AgarAgar, a gelatinous substance derived from red algae, has a number of commercial uses. It is a good medium on which to grow bacteria and fungi, as most microorganisms cannot digest agar.
AlginatesAlginic acid, or alginate, is extracted from brown algae. Its uses range from gelling agents in food, to medical dressings. Alginic acid also has been used in the field of biotechnology as a biocompatible medium for cell encapsulation and cell immobilization. Molecular cuisine is also a user of the substance for its gelling properties, by which it becomes a delivery vehicle for flavours. Between 100,000 and 170,000 wet tons of Macrocystis are harvested annually in New Mexico for alginate extraction and abalone feed.
Energy sourceTo 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 is estimated to occur by 2025.An Outlook on Microalgal Biofuels, René H. Wijffels and Maria J. Barbosa, Science 13 August 2010: 329 (5993), 796–799.
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: Downloadable Google Books .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. 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.
NutritionNaturally 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. In addition, commercially cultivated microalgae, including both algae and cyanobacteria, are marketed as nutritional supplements, such as spirulina, Chlorella and the vitamin-C supplement from 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; Ireland, dulse; Chile, cochayuyo. 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 salad ingredients in Scotland, Ireland, Greenland, and Iceland. 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. 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). 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. Algae have 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).
- Sewage can be treated with algae, 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. Hydromentia Water Treatment Technologies
PolymersVarious 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. Several companies have begun to produce algae polymers commercially, including for use in flip-flops and in surf boards.
BioremediationThe alga Stichococcus bacillaris has been seen to colonize silicone resins used at archaeological sites; biodegrading the synthetic substance.
PigmentsThe natural pigments ( carotenoids and chlorophylls) produced by algae can be used as alternatives to chemical dyes and coloring agents. 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 alga populations in sea water samples.
Stabilizing substancesCarrageenan, from the red alga Chondrus crispus, is used as a stabilizer in milk products.
- Toxoid - anatoxin
- Marimo algae
- Iron fertilization
- Fritsch, F.E. (1935/1945). The Structure and Reproduction of the Algae. I. and II. Cambridge, England: Cambridge University Press
- van den Hoek, C., D.G. Mann, and H.M. Jahns (1995). Algae: an introduction to phycology. Cambridge University Press (623 pp).
- Smith, G.M. (1938). Cryptogamic Botany, vol. 1. McGraw-Hill, New York.
Regional;Britain and Ireland
- – a database of all algal names including images, nomenclature, taxonomy, distribution, bibliography, uses, extracts
- Algae – Cell Centered Database
- Algae: Protists with Chloroplasts
- Algae glossary (Australian Biological Resources Study).
- EnAlgae http://www.enalgae.eu