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{{redirect|Symbiology|use of things that represent other things by association, resemblance, or convention|Symbology}}{{About|the biological phenomenon|other uses|Symbiosis (disambiguation)}}File:Common clownfish curves dnsmpl.jpg|thumb|upright=1.7|In a cleaning symbiosis, the clownfish feeds on small invertebrates that otherwise have potential to harm the sea anemone, and the fecal matter from the clownfish provides nutrients to the sea anemone. The clownfish is protected from predators by the anemone's stinging cells, to which the clownfish is immune. The clownfish emits a high pitched sound that deters butterfly fish, which would otherwise eat the anemone, making the relationship appear (Mutualism (biology)|mutualistic]].WEB, Miller, Allie, Intricate Relationship Allows the Other to Flourish: the Sea Anemone and the Clownfish,weblink AskNature, The Biomimicry Institute, 15 February 2015, )Symbiosis (from Greek συμβίωσις "living together", from σύν "together" and βίωσις "living"){{LSJ|sumbi/wsis|συμβίωσις}}, {{LSJ|su/n|σύν}}, {{LSJ|bi/wsis|βίωσις|ref}} is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasitic. The organisms, each termed a symbiont, may be of the same or of different species. In 1879, Heinrich Anton de Bary defined it as "the living together of unlike organisms". The term was subject to a century-long debate about whether it should specifically denote mutualism, as in lichens; biologists have now abandoned that restriction.Symbiosis can be obligatory, which means that one or both of the symbionts entirely depend on each other for survival, or facultative (optional) when they can generally live independently.Symbiosis is also classified by physical attachment; symbiosis in which the organisms have bodily union is called conjunctive symbiosis, and symbiosis in which they are not in union is called disjunctive symbiosis."symbiosis." Dorland's Illustrated Medical Dictionary. Philadelphia: Elsevier Health Sciences, 2007. Credo Reference. Web. 17 September 2012When one organism lives on the surface of another, such as head lice on humans, it is called ectosymbiosis; when one partner lives inside the tissues of another, such as Symbiodinium within coral, it is termed endosymbiosis.{{Harvnb|Moran|2006}}{{Harvnb|Paracer|Ahmadjian|2000|p=12}}

Definition

(File:Symbiotic relationships diagram.svg|thumb|upright=1.4|Diagram of the six possible types of symbiotic relationship, from mutual benefit to mutual harm.)The definition of symbiosis was a matter of debate for 130 years.{{Citation |title= Symbiosis: 'Living together' in chaos|last= Martin|first= Bradford D.|last2= Schwab|first2= Ernest|year= 2012|journal= Studies in the History of Biology|volume= 4|issue= 4|pages= 7–25}} In 1877, Albert Bernhard Frank used the term symbiosis to describe the mutualistic relationship in lichens.{{OED|symbiosis}} In 1879, the German mycologist Heinrich Anton de Bary defined it as "the living together of unlike organisms".{{Harvnb|Wilkinson|2001}}{{Harvnb|Douglas|1994|p=1}} The definition has varied among scientists, with some advocating that it should only refer to persistent mutualisms, while others thought it should apply to all persistent biological interactions, in other words mutualisms, commensalism, or parasitism, but excluding brief interactions such as predation.{{harvnb|Douglas|2010|pp=5–12}} Current biology and ecology textbooks use the latter "de Bary" definition, or an even broader one where symbiosis means all interspecific interactions; the restrictive definition where symbiosis means only mutualism is no longer used.{{Citation |title= Current usage of symbiosis and associated terminology|last= Martin|first= Bradford D.|last2= Schwab|first2= Ernest|year= 2013|journal= International Journal of Biology|volume= 5|issue= 1|pages= 32–45|doi=10.5539/ijb.v5n1p32}}In 1949, Edward Haskell proposed an integrative approach, proposing a classification of "co-actions",Haskell, E. F. (1949). A clarification of social science. Main Currents in Modern Thought 7: 45–51. later adopted by biologists as "interactions".Burkholder, P. R. (1952) Cooperation and Conflict among Primitive Organisms. American Scientist, 40, 601-631. link.Bronstein, J. L. (2015). The study of mutualism. In: Bronstein, J. L. (ed.). Mutualism. Oxford University Press, Oxford. link.JOURNAL, 10.1371/journal.pbio.2000891, 27732591, 5061325, Orienting the Interaction Compass: Resource Availability as a Major Driver of Context Dependence, PLOS Biology, 14, 10, e2000891, 2016, Pringle, Elizabeth G., Biological interactions can involve individuals of the same species (intraspecific interactions) or individuals of different species (interspecific interactions). These can be further classified by either the mechanism of the interaction or the strength, duration and direction of their effects.JOURNAL, Wootton, J.T., Emmerson, M., Measurement of Interaction Strength in Nature, Annual Review of Ecology, Evolution, and Systematics, 36, 419–44, 2005, 30033811, 10.1146/annurev.ecolsys.36.091704.175535,

Obligate versus facultative

Relationships can be obligate, meaning that one or both of the symbionts entirely depend on each other for survival. For example, in lichens, which consist of fungal and photosynthetic symbionts, the fungal partners cannot live on their own.{{Harvnb|Isaac|1992|p=266}}{{Harvnb|Saffo|1993}}{{harvnb|Douglas|2010|p=4}} The algal or cyanobacterial symbionts in lichens, such as Trentepohlia, can generally live independently, and their symbiosis is, therefore, facultative (optional).JOURNAL, Muggia, Lucia, Vancurova, Lucie, Škaloud, Pavel, Peksa, Ondrej, Wedin, Mats, Grube, Martin, The symbiotic playground of lichen thalli - a highly flexible photobiont association in rock-inhabiting lichens, FEMS Microbiology Ecology, 85, 2, 2013, 10.1111/1574-6941.12120, 23530593, 313–323,

Physical interaction

File:An alder root nodule gall.JPG|right|thumb|Alder tree root nodule houses endosymbiotic nitrogen-fixing bacterianitrogen-fixing bacteria{{See also|Root microbiome}}Endosymbiosis is any symbiotic relationship in which one symbiont lives within the tissues of the other, either within the cells or extracellularly.{{Harvnb|Sapp|1994|p=142}} Examples include diverse microbiomes, rhizobia, nitrogen-fixing bacteria that live in root nodules on legume roots; actinomycete, nitrogen-fixing bacteria such as Frankia, which live in alder root nodules; single-celled algae inside reef-building corals; and bacterial endosymbionts that provide essential nutrients to about 10%–15% of insects.{{citation needed|date=March 2017}}Ectosymbiosis is any symbiotic relationship in which the symbiont lives on the body surface of the host, including the inner surface of the digestive tract or the ducts of exocrine glands.{{Harvnb|Nardon|Charles|2002}} Examples of this include ectoparasites such as lice; commensal ectosymbionts such as the barnacles, which attach themselves to the jaw of baleen whales; and mutualist ectosymbionts such as cleaner fish.File:Hirschkampf.jpg|right|thumb|Male-male interference competition in red deerred deer

Competition

Competition can be defined as an interaction between organisms or species, in which the fitness of one is lowered by the presence of another. Limited supply of at least one resource (such as food, water, and territory) used by both usually facilitates this type of interaction, although the competition may also exist over other 'amenities', such as females for reproduction (in case of male organisms of the same species).Begon, M., J.L. Harper and C.R. Townsend. 1996. Ecology: individuals, populations, and communities, Third Edition. Blackwell Science Ltd., Cambridge, Massachusetts, USA.

Mutualism

File:Calcinus laevimanus hermit crab with Calliactis sea anemone. 2 frames in one.jpg|thumb|300pxMutualism or interspecies reciprocal altruism is a long-term relationship between individuals of different species where both individuals benefit.{{Harvnb|Paracer|Ahmadjian|2000|p=6}} Mutualistic relationships may be either obligate for both species, obligate for one but facultative for the other, or facultative for both.File:Bryolith (Banc d'Arguin, Mauritania).jpg|thumb|upright|Bryoliths document a mutualistic symbiosis between a hermit crab and encrusting bryozoansbryozoansA large percentage of herbivores have mutualistic gut flora to help them digest plant matter, which is more difficult to digest than animal prey. This gut flora is made up of cellulose-digesting protozoans or bacteria living in the herbivores' intestines."symbiosis." The Columbia Encyclopedia. New York: Columbia University Press, 2008. Credo Reference. Web. 17 September 2012. Coral reefs are the result of mutualisms between coral organisms and various types of algae which live inside them.{{Harvnb|Toller|Rowan|Knowlton|2001}} Most land plants and land ecosystems rely on mutualisms between the plants, which fix carbon from the air, and mycorrhyzal fungi, which help in extracting water and minerals from the ground.{{Harvnb|Harrison|2005}}An example of mutualism is the relationship between the ocellaris clownfish that dwell among the tentacles of Ritteri sea anemones. The territorial fish protects the anemone from anemone-eating fish, and in turn the stinging tentacles of the anemone protect the clownfish from its predators. A special mucus on the clownfish protects it from the stinging tentacles.{{Harvnb|Lee|2003}}A further example is the goby, a fish which sometimes lives together with a shrimp. The shrimp digs and cleans up a burrow in the sand in which both the shrimp and the goby fish live. The shrimp is almost blind, leaving it vulnerable to predators when outside its burrow. In case of danger, the goby touches the shrimp with its tail to warn it. When that happens both the shrimp and goby quickly retreat into the burrow.{{Harvnb|Facey|Helfman|Collette|1997}} Different species of gobies (Elacatinus spp.) also clean up ectoparasites in other fish, possibly another kind of mutualism.JOURNAL, M.C. Soares, I.M. Côté, S.C. Cardoso & R.Bshary, August 2008, The cleaning goby mutualism: a system without punishment, partner switching or tactile stimulation, Journal of Zoology, 276, 3, 306–312, 10.1111/j.1469-7998.2008.00489.x,weblink A non-obligate symbiosis is seen in encrusting bryozoans and hermit crabs. The bryozoan colony (Acanthodesia commensale) develops a cirumrotatory growth and offers the crab (Pseudopagurus granulimanus) a helicospiral-tubular extension of its living chamber that initially was situated within a gastropod shell.JOURNAL, Klicpera, A, PD Taylor, H Westphal, Bryoliths constructed by bryozoans in symbiotic associations with hermit crabs in a tropical heterozoan carbonate system, Golfe d'Arguin, Mauritania, Mar Biodivers, 1 Dec 2013, 43, 4, 429–444, 10.1007/s12526-013-0173-4, 1867-1616, Many types of tropical and sub-tropical ants have evolved very complex relationships with certain tree species.Piper, Ross (2007), Extraordinary Animals: An Encyclopedia of Curious and Unusual Animals, Greenwood Press.

Endosymbiosis

{{further|Endosymbiont}}In endosymbiosis, the host cell lacks some of the nutrients which the endosymbiont provides. As a result, the host favors endosymbiont's growth processes within itself by producing some specialized cells. These cells affect the genetic composition of the host in order to regulate the increasing population of the endosymbionts and ensure that these genetic changes are passed onto the offspring via vertical transmission (heredity).BOOK, Latorre, A., The role of symbiosis in eukaryotic evolution. Origins and evolution of life – An astrobiological perspective, 2011, 326–339, Durban, A., Moya, A., Pereto, J., A spectacular example of obligate mutualism is the relationship between the siboglinid tube worms and symbiotic bacteria that live at hydrothermal vents and cold seeps. The worm has no digestive tract and is wholly reliant on its internal symbionts for nutrition. The bacteria oxidize either hydrogen sulfide or methane, which the host supplies to them. These worms were discovered in the late 1980s at the hydrothermal vents near the Galapagos Islands and have since been found at deep-sea hydrothermal vents and cold seeps in all of the world's oceans.{{harvnb|Cordes|Arthur|Shea|Arvidson|2005}}As the endosymbiont adapts to the host's lifestyle, the endosymbiont changes dramatically. There is a drastic reduction in its genome size, as many genes are lost during the process of metabolism, and DNA repair and recombination, while important genes participating in the DNA-to-RNA transcription, protein translation and DNA/RNA replication are retained. The decrease in genome size is due to loss of protein coding genes and not due to lessening of inter-genic regions or open reading frame (ORF) size. Species that are naturally evolving and contain reduced sizes of genes can be accounted for an increased number of noticeable differences between them, thereby leading to changes in their evolutionary rates. When endosymbiotic bacteria related with insects are passed on to the offspring strictly via vertical genetic transmission, intracellular bacteria go across many hurdles during the process, resulting in the decrease in effective population sizes, as compared to the free-living bacteria. The incapability of the endosymbiotic bacteria to reinstate their wild type phenotype via a recombination process is called Muller's ratchet phenomenon. Muller's ratchet phenomenon, together with less effective population sizes, leads to an accretion of deleterious mutations in the non-essential genes of the intracellular bacteria.JOURNAL, Moran, N. A., Accelerated evolution and Muller's ratchet in endosymbiotic bacteria, Proceedings of the National Academy of Sciences of the United States of America, 1996, 93, 7, 2873–2878, 10.1073/pnas.93.7.2873, 8610134, 39726, This can be due to lack of selection mechanisms prevailing in the relatively "rich" host environment.JOURNAL, Andersson, Siv G.E, Kurland, Charles G, Reductive evolution of resident genomes, Trends in Microbiology, 1998, 6, 7, 263–8, 10.1016/S0966-842X(98)01312-2, 9717214, JOURNAL, Wernegreen, J.J., Genome evolution in bacterial endosymbionts of insects, Nature Reviews Genetics, 2002, 3, 850–861, 10.1038/nrg931, 12415315, 11,

Commensalism

File:Fly June 2008-2.jpg|right|140px|thumb|Commensal mites travelling (phoresy) on a fly (Pseudolynchia canariensisPseudolynchia canariensisCommensalism describes a relationship between two living organisms where one benefits and the other is not significantly harmed or helped. It is derived from the English word (wikt:commensal|commensal), used of human social interaction. It derives from a medieval Latin word meaning sharing food, formed from com- (with) and mensa (table).{{Harvnb|Nair|2005}}Commensal relationships may involve one organism using another for transportation (phoresy) or for housing (inquilinism), or it may also involve one organism using something another created, after its death (metabiosis). Examples of metabiosis are hermit crabs using gastropod shells to protect their bodies, and spiders building their webs on plants.

Parasitism

File:Taenia solium tapeworm scolex with its four suckers and two rows of hooks 5262 lores.jpg|thumb|Head (scolex) of tapeworm Taenia solium is adapted to parasitism with hooks and suckers to attach to its host.]] In a parasitic relationship, the parasite benefits while the host is harmed.{{Harvnb|Paracer|Ahmadjian|2000|p=7}} Parasitism takes many forms, from endoparasites that live within the host's body to ectoparasites and parasitic castrators that live on its surface and micropredators like mosquitoes that visit intermittently. Parasitism is an extremely successful mode of life; as many as half of all animals have at least one parasitic phase in their life cycles, and it is also frequent in plants and fungi. Moreover, almost all free-living animal species are hosts to parasites, often of more than one species.{{citation needed|date=February 2018}}

Mimicry

Mimicry is a form of symbiosis in which a species adopts distinct characteristics of another species to alter its relationship dynamic with the species being mimicked, to its own advantage. Among the many types of mimicry are Batesian and Müllerian, the first involving one-sided exploitation, the second providing mutual benefit. Batesian mimicry is an exploitative three-party interaction where one species, the mimic, has evolved to mimic another, the model, to deceive a third, the dupe. In terms of signalling theory, the mimic and model have evolved to send a signal; the dupe has evolved to receive it from the model. This is to the advantage of the mimic but to the detriment of both the model, whose protective signals are effectively weakened, and of the dupe, which is deprived of an edible prey. For example, a wasp is a strongly-defended model, which signals with its conspicuous black and yellow coloration that it is an unprofitable prey to predators such as birds which hunt by sight; many hoverflies are Batesian mimics of wasps, and any bird that avoids these hoverflies is a dupe.JOURNAL, Vane-Wright, R. I., 1976, A unified classification of mimetic resemblances, Biological Journal of the Linnean Society, 8, 25–56, 10.1111/j.1095-8312.1976.tb00240.x, JOURNAL, Bates, Henry Walter, Henry Walter Bates, 1861, Contributions to an insect fauna of the Amazon valley. Lepidoptera: Heliconidae,weblink Transactions of the Linnean Society of London, Transactions of the Linnean Society, 23, 3, 495–566, 10.1111/j.1096-3642.1860.tb00146.x, ; Reprint: JOURNAL, Bates, Henry Walter, 1981, Contributions to an insect fauna of the Amazon valley (Lepidoptera: Heliconidae), Biological Journal of the Linnean Society, 16, 1, 41–54, 10.1111/j.1095-8312.1981.tb01842.x, In contrast, Müllerian mimicry is mutually beneficial as all participants are both models and mimics.JOURNAL, Müller, Fritz, Fritz Müller, 1878, Ueber die Vortheile der Mimicry bei Schmetterlingen, Zoologischer Anzeiger, 1, 54–55, JOURNAL, Müller, Fritz, Fritz Müller, 1879, Ituna and Thyridia; a remarkable case of mimicry in butterflies. (R. Meldola translation), Proclamations of the Entomological Society of London, 1879, 20–29, For example, different species of bumblebee mimic each other, with similar warning coloration in combinations of black, white, red, and yellow, and all of them benefit from the relationship.JOURNAL, Mallet, James, James Mallet, Causes and consequences of a lack of coevolution in Mullerian mimicry, Evolutionary Ecology, 2001, 13, 7–8, 777–806, 10.1023/a:1011060330515, 10.1.1.508.2755,

Amensalism

File:Black Walnut middle.JPG|right|thumb|The black walnut secretes a chemical from its roots that harms neighboring plants, an example of antagonism.]]Amensalism is an asymmetric interaction where one species is harmed or killed by the other, and one is unaffected by the other.Toepfer, G. "Amensalism". In: BioConcepts. link.BOOK, Willey, Joanne M., Sherwood, Linda M., Woolverton, Cristopher J., 2013, Prescott's Microbiology, 9th, 713–738, 978-0-07-751066-4, There are two types of amensalism, competition and antagonism (or antibiosis). Competition is where a larger or stronger organism deprives a smaller or weaker one from a resource. Antagonism occurs when one organism is damaged or killed by another through a chemical secretion. An example of competition is a sapling growing under the shadow of a mature tree. The mature tree can rob the sapling of necessary sunlight and, if the mature tree is very large, it can take up rainwater and deplete soil nutrients. Throughout the process, the mature tree is unaffected by the sapling. Indeed, if the sapling dies, the mature tree gains nutrients from the decaying sapling. An example of antagonism is Juglans nigra (black walnut), secreting juglone, a substance which destroys many herbaceous plants within its root zone.The Editors of Encyclopædia Britannica. (n.d.). Amensalism (biology). Retrieved September 30, 2014, fromweblink clear case of amensalism is where sheep or cattle trample grass. Whilst the presence of the grass causes negligible detrimental effects to the animal's hoof, the grass suffers from being crushed.{{citation needed|date=February 2018}} Amensalism is often used to describe strongly asymmetrical competitive interactions, such as has been observed between the Spanish ibex and weevils of the genus Timarcha which feed upon the same type of shrub. Whilst the presence of the weevil has almost no influence on food availability, the presence of ibex has an enormous detrimental effect on weevil numbers, as they consume significant quantities of plant matter and incidentally ingest the weevils upon it.JOURNAL, Gómez, José M., González-Megías, Adela, 2002, Asymmetrical interactions between ungulates and phytophagous insects: Being different matters, Ecology, 83, 1, 203–11, 10.1890/0012-9658(2002)083[0203:AIBUAP]2.0.CO;2,

Cleaning symbiosis

Cleaning symbiosis is an association between individuals of two species, where one (the cleaner) removes and eats parasites and other materials from the surface of the other (the client).JOURNAL, Losey, G.S., The Ecological Importance of Cleaning Symbiosis, Copeia, 1972, 4, 1972, 820–833, 10.2307/1442741, 1442741, It is putatively mutually beneficial, but biologists have long debated whether it is mutual selfishness, or simply exploitative. Cleaning symbiosis is well-known among marine fish, where some small species of cleaner fish, notably wrasses but also species in other genera, are specialised to feed almost exclusively by cleaning larger fish and other marine animals.JOURNAL, Poulin, Robert, Robert Poulin (zoologist), Grutter, A. S., 1996, Cleaning symbiosis: proximate and adaptive explanations, BioScience, 46, 7, 512–517,weblink 10.2307/1312929, 1312929,

Co-evolution

File:Common jassid nymphs and ants02.jpg|thumb|upright|Leafhoppers protected by meat antmeat ant{{further|Co-evolution}}Symbiosis is increasingly recognized as an important selective force behind evolution;{{Harvnb|Wernegreen|2004}} many species have a long history of interdependent co-evolution.{{Harvnb|Paracer|Ahmadjian|2000|pp=3–4}}

Symbiogenesis

Eukaryotes (plants, animals, fungi, and protists) developed by symbiogenesis from a symbiosis between bacteria and archaea.{{Harvnb|Brinkman|Blanchard|Cherkasov|Av-gay|2002}}{{Harvnb|Golding|Gupta|1995}} Evidence for this includes the fact that mitochondria and chloroplasts divide independently of the cell, and the observation that some organelles seem to have their own genome."Symbiosis". Bloomsbury Guide to Human Thought. London: Bloomsbury Publishing Ltd, 1993. Credo Reference. Web. 17 September 2012.The biologist Lynn Margulis, famous for her work on endosymbiosis, contended that symbiosis is a major driving force behind evolution. She considered Darwin's notion of evolution, driven by competition, to be incomplete and claimed that evolution is strongly based on co-operation, interaction, and mutual dependence among organisms. According to Margulis and her son Dorion Sagan, "Life did not take over the globe by combat, but by networking."{{harvnb|Sagan|Margulis|1986}}

Co-evolutionary relationships

Mycorrhizas

About 80% of vascular plants worldwide form symbiotic relationships with fungi, in particular in arbuscular mycorrhizas.{{Citation | author= Schüßler, A. | year=2001 | title=A new fungal phylum, the Glomeromycota: phylogeny and evolution | journal=Mycol. Res. | volume=105 | issue=12 | pages=1413–1421 | url=http://journals.cambridge.org/action/displayAbstract?fromPage=online&aid=95091 | doi= 10.1017/S0953756201005196 | postscript= .|display-authors=etal}}File:Hummingbird hawkmoth a.jpg|upright|thumb|Pollination is a mutualism between flowering plantflowering plant

Pollination

File:Ficus plant.jpg|thumb|left|A fig is pollinated by the fig wasp, Blastophaga psenesBlastophaga psenes{{further|Entomophily|Ornithophily|Reproductive coevolution in Ficus}}Flowering plants and the animals that pollinate them have co-evolved. Many plants that are pollinated by insects (in entomophily), bats, or birds (in ornithophily) have highly specialized flowers modified to promote pollination by a specific pollinator that is correspondingly adapted. The first flowering plants in the fossil record had relatively simple flowers. Adaptive speciation quickly gave rise to many diverse groups of plants, and, at the same time, corresponding speciation occurred in certain insect groups. Some groups of plants developed nectar and large sticky pollen, while insects evolved more specialized morphologies to access and collect these rich food sources. In some taxa of plants and insects, the relationship has become dependent,{{Harvnb|Harrison|2002}} where the plant species can only be pollinated by one species of insect.{{Harvnb|Danforth|Ascher|1997}}File:Ant - Pseudomyrmex species, on Bull Thorn Acacia (Acacia cornigera) with Beltian bodies, Caves Branch Jungle Lodge, Belmopan, Belize - 8505045055.jpg|thumb|Pseudomyrmex ant on bull thorn acacia (Vachellia cornigeraVachellia cornigera

Acacia ants and acacias

The acacia ant (Pseudomyrmex ferruginea) is an obligate plant ant that protects at least five species of "Acacia" (Vachellia){{efn|The acacia ant protects at least 5 species of "Acacia", now all renamed to Vachellia: V. chiapensis, V. collinsii, V. cornigera, V. hindsii and V. sphaerocephala.}} from preying insects and from other plants competing for sunlight, and the tree provides nourishment and shelter for the ant and its larvae.BOOK, Hölldobler, Bert, Wilson, Edward O., The ants, Harvard University Press, 1990,weblink 978-0-674-04075-5, 532–533, WEB, National Geographic, Acacia Ant Video,weblink yes,weblink" title="web.archive.org/web/20071107085438weblink">weblink 2007-11-07,

See also

Notes

{{notelist}}

References

{{reflist|30em}}

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