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Symbiosis

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This article is about the biological phenomenon. For other uses, seeSymbiosis (disambiguation).

Symbiosis(Ancient Greekσυμβίωσιςsymbíōsis:living with, companionship <σύνsýn:together; andβίωσιςbíōsis:living)[2]is any type of a close and long-termbiological interaction,between twoorganismsof differentspecies.The two organisms, termedsymbionts,can be either in amutualistic,acommensalistic,or aparasiticrelationship.[3]In 1879,Heinrich Anton de Barydefined symbiosis as "the living together of unlike organisms".

In acleaning symbiosis,theclownfishfeeds on small invertebrates, that otherwise have potential to harm thesea 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 relationship is therefore classified asmutualistic.[1]

The term is sometimes more exclusively used in a restricted, mutualistic sense, where both symbionts contribute to each other's subsistence.[3]

Symbiosis can beobligatory,which means that one, or both of the symbionts depend on each other for survival, orfacultative(optional), when they can also subsist independently.

Symbiosis is also classified by physical attachment. Symbionts forming a single body live inconjunctivesymbiosis, while all other arrangements are calleddisjunctivesymbiosis.[4]When one organism lives on the surface of another, such ashead liceon humans, it is calledectosymbiosis;when one partner lives inside the tissues of another, such asSymbiodiniumwithincoral,it is termedendosymbiosis.[5][6]

Definition

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Diagram of the six possible types of symbiotic relationship, from mutual benefit to mutual harm.

The definition ofsymbiosiswas a matter of debate for 130 years.[7]In 1877,Albert Bernhard Frankused the termsymbiosisto describe the mutualistic relationship inlichens.[8][9]In 1878, the German mycologistHeinrich Anton de Barydefined it as "the living together of unlike organisms".[10][11][12]The definition has varied among scientists, with some advocating that it should only refer to persistentmutualisms,while others thought it should apply to all persistent biological interactions (in other words, to mutualism,commensalism,andparasitism,but excluding brief interactions such aspredation). In the 21st century, the latter has become the definition widely accepted by biologists.[13]

In 1949,Edward Haskellproposed an integrative approach with a classification of "co-actions",[14]later adopted by biologists as "interactions".[15][16][17][18]

Types

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Obligate versus facultative

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Relationships can be obligate, meaning that one or both of the symbionts entirely depend on each other for survival. For example, inlichens,which consist of fungal andphotosyntheticsymbionts, the fungal partners cannot live on their own.[11][19][20][21]The algal or cyanobacterial symbionts in lichens, such asTrentepohlia,can generally live independently, and their part of the relationship is therefore described as facultative (optional), or non-obligate.[22]When one of the participants in a symbiotic relationship is capable of photosynthesis, as with lichens, it is called photosymbiosis.[23][24]

Ectosymbiosis versus endosymbiosis

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Alder tree root nodule houses endosymbioticnitrogen-fi xing bacteria.
Main article:Ectosymbiosis
Further information:Endosymbiont

Ectosymbiosisis any symbiotic relationship in which the symbiont lives on the body surface of thehost,including the inner surface of thedigestivetract or the ducts ofexocrine glands.[6][25]Examples of this includeectoparasitessuch aslice;commensalectosymbionts such as thebarnacles,which attach themselves to the jaw ofbaleen whales;and mutualist ectosymbionts such ascleaner fish.

Contrastingly,endosymbiosisis any symbiotic relationship in which one symbiont lives within the tissues of the other, either within the cells or extracellularly.[6][26]Examples include diversemicrobiomes:rhizobia,nitrogen-fi xing bacteriathat live inroot nodulesonlegumeroots;actinomycetes,nitrogen-fi xing bacteria such asFrankia,which live inalderroot nodules; single-celledalgaeinside reef-buildingcorals;and bacterialendosymbiontsthat provide essential nutrients to about 10%–15% of insects.[27]

In endosymbiosis, the host cell lacks some of the nutrients which theendosymbiontprovides. 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 viavertical transmission(heredity).[28]

As the endosymbiont adapts to the host's lifestyle, the endosymbiont changes dramatically. There is a drastic reduction in itsgenomesize, as many genes are lost during the process ofmetabolism,andDNArepair and recombination, while important genes participating in the DNA-to-RNAtranscription,proteintranslationand 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 oropen 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 typephenotypevia a recombination process is calledMuller's ratchetphenomenon. Muller's ratchet phenomenon, together with less effective population sizes, leads to an accretion of deleteriousmutationsin the non-essential genes of the intracellular bacteria.[29]This can be due to lack ofselectionmechanisms prevailing in the relatively "rich" host environment.[30][31]

Competition

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Main article:Competition (biology)

Competition can be defined as an interaction betweenorganismsor species, in which thefitnessof one is lowered by the presence of another.[32]Limitedsupply of at least one resource (such asfood,water,andterritory) used by both usually facilitates this type of interaction, although the competition can also be for other resources.[33]

Amensalism

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Theblack walnutsecretes a chemical from its roots that harms neighboring plants, an example ofantagonism.

Amensalism is a non-symbiotic, asymmetric interaction where one species is harmed or killed by the other, and one is unaffected by the other.[34][35]There are two types of amensalism, competition andantagonism(or antibiosis). Competition is where a larger or stronger organism deprives a smaller or weaker one of 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 thesaplingof 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 isJuglans nigra(black walnut), secretingjuglone,a substance which destroys many herbaceous plants within its root zone.[36]

The termamensalismis often used to describe strongly asymmetrical competitive interactions, such as between theSpanish ibexandweevilsof the genusTimarchawhich 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.[37]

Commensalism

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Main article:Commensalism
Commensalmites travelling (phoresy) on a fly (Pseudolynchia canariensis)

Commensalism 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 wordcommensal,used of humansocial interaction.It derives from a medieval Latin word meaning sharing food, formed fromcom-(with) andmensa(table).[38][39]

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 arehermit crabsusinggastropodshells to protect their bodies, and spiders building their webs onplants.

Mutualism

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Main article:Mutualism (biology)
Hermit crab,Calcinus laevimanus,with sea anemone

Mutualism or interspeciesreciprocal altruismis a long-term relationship between individuals of differentspecieswhere both individuals benefit.[38]Mutualistic relationships may be either obligate for both species, obligate for one but facultative for the other, or facultative for both.

Bryolithsdocument a mutualistic symbiosis between ahermit craband encrustingbryozoans.

Manyherbivoreshave mutualisticgut florato help them digest plant matter, which is more difficult to digest than animal prey.[5]This gut flora comprises cellulose-digestingprotozoansor bacteria living in the herbivores' intestines.[40]Coralreefs result from mutualism between coral organisms and various algae living inside them.[41]Most land plants and land ecosystems rely on mutualism between the plants, whichfixcarbon from the air, andmycorrhyzalfungi, which help in extracting water and minerals from the ground.[42]

An example of mutualism is the relationship between theocellaris clownfishthat dwell among thetentaclesofRitteri sea anemones.The territorial fish protects the anemone from anemone-eating fish, and in turn, the anemone stinging tentacles protect the clownfish from itspredators.A specialmucuson the clownfish protects it from the stinging tentacles.[43]

A further example is thegoby,a fish which sometimes lives together with ashrimp.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, and both quickly retreat into the burrow.[44]Different species of gobies (Elacatinusspp.) alsoclean up ectoparasitesin other fish, possibly another kind of mutualism.[45]

A spectacular example of obligate mutualism is the relationship between thesiboglinidtube wormsand symbioticbacteriathat live athydrothermal ventsandcold seeps.The worm has no digestive tract and is wholly reliant on its internal symbionts for nutrition. The bacteria oxidize eitherhydrogen sulfideor 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 atdeep-seahydrothermal vents and cold seeps in all of the world's oceans.[46]

Mutualism improves both organism's competitive ability and will outcompete organisms of the same species that lack the symbiont.[47]

A facultative symbiosis is seen in encrustingbryozoansandhermit 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.[48]

Parasitism

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Main article:Parasitism
Head (scolex) oftapewormTaenia soliumis adapted toparasitismwith hooks and suckers to attach to itshost.

In a parasitic relationship, the parasite benefits while the host is harmed.[49]Parasitism takes many forms, fromendoparasitesthat live within the host's body toectoparasitesandparasitic castratorsthat live on its surface andmicropredatorslike mosquitoes that visit intermittently. Parasitism is an extremely successful mode of life; about 40% of all animal species are parasites, and the average mammal species is host to 4 nematodes, 2 cestodes, and 2 trematodes.[50]

Mimicry

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Main article: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 mimicryis an exploitative three-party interaction where one species, the mimic, has evolved to mimic another, the model, todeceivea third, the dupe. In terms ofsignalling 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.[51][52]In contrast,Müllerian mimicryis mutually beneficial as all participants are both models and mimics.[53][54]For example, different species ofbumblebeemimic each other, with similar warning coloration in combinations of black, white, red, and yellow, and all of them benefit from the relationship. [55]

Cleaning symbiosis

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Main article:Cleaning symbiosis

Cleaning symbiosisis 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).[56]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 ofcleaner fish– notablywrasses,but also species in other genera – are specialized to feed almost exclusively by cleaning larger fish and other marine animals.[57]In a supreme situation, the host species (fish or marine life) will display itself at a designated station deemed the "cleaning station".[58]

Cleaner fish play an essential role in the reduction of parasitism on marine animals. Some shark species participate in cleaning symbiosis, where cleaner fish remove ectoparasites from the body of the shark.[59]A study by Raymond Keyes addresses the atypical behavior of a few shark species when exposed to cleaner fish. In this experiment, cleaner wrasse(Labroides dimidiatus)and various shark species were placed in a tank together and observed. The different shark species exhibited different responses and behaviors around the wrasse. For example, Atlantic and Pacific lemon sharks consistently react to the wrasse fish in a fascinating way. During the interaction, the shark remains passive and the wrasse swims to it. It begins to scan the shark's body, sometimes stopping to inspect specific areas. Commonly, the wrasse would inspect the gills, labial regions, and skin. When the wrasse makes its way to the mouth of the shark, the shark often ceases breathing for up to two and a half minutes so that the fish is able to scan the mouth. Then, the fish passes further into the mouth to examine the gills, specifically the buccopharyngeal area, which typically holds the most parasites. When the shark begins to close its mouth, the wrasse finishes its examination and goes elsewhere. Male bull sharks exhibit slightly different behavior at cleaning stations: as the shark swims into a colony of wrasse fish, it drastically slows its speed to allow the cleaners to do their job. After approximately one minute, the shark returns to normal swimming speed.[59]

Role in evolution

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Leafhoppersprotected bymeat ants
Further information:Co-evolution

Symbiosis is increasingly recognized as an important selective force behind evolution;[5][60]many species have a long history of interdependentco-evolution.[61]

Although symbiosis was once discounted as an anecdotal evolutionary phenomenon, evidence is now overwhelming that obligate or facultative associations among microorganisms and between microorganisms and multicellular hosts had crucial consequences in many landmark events in evolution and in the generation of phenotypic diversity and complex phenotypes able to colonise new environments.[62]

Hologenome development and evolution

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Evolution originated from changes in development where variations within species are selected for or against because of the symbionts involved.[63]The hologenome theory relates to the holobiont and symbionts genome together as a whole.[64]Microbes live everywhere in and on every multicellular organism.[65]Many organisms rely on their symbionts in order to develop properly, this is known as co-development. In cases of co-development the symbionts send signals to their host which determine developmental processes. Co-development is commonly seen in both arthropods and vertebrates.[63]

Symbiogenesis

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Main article:Symbiogenesis

One hypothesis for the origin of the nucleus ineukaryotes(plants, animals, fungi, andprotists) is that it developed from asymbiogenesisbetween bacteria and archaea.[5][66][67]It is hypothesized that the symbiosis originated when ancient archaea, similar to modern methanogenic archaea, invaded and lived within bacteria similar to modern myxobacteria, eventually forming the early nucleus. This theory is analogous to the accepted theory for the origin of eukaryotic mitochondria and chloroplasts, which are thought to have developed from a similar endosymbiotic relationship between proto-eukaryotes and aerobic bacteria.[68]Evidence for this includes the fact thatmitochondriaandchloroplastsdivide independently of the cell, and that these organelles have their own genome.[69]

The biologistLynn Margulis,famous for her work onendosymbiosis,contended that symbiosis is a major driving force behindevolution.She consideredDarwin's notion of evolution, driven by competition, to be incomplete and claimed that evolution is strongly based onco-operation,interaction,andmutual dependenceamong organisms. According to Margulis and her sonDorion Sagan,"Lifedid not take over theglobebycombat,but bynetworking."[70]

Major examples of co-evolutionary relationships

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Mycorrhiza

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About 80% ofvascular plantsworldwide form symbiotic relationships with fungi, in particular inarbuscular mycorrhizas.[71]

Pollination is a mutualism betweenflowering plantsand their animal pollinators.

Pollination

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Afigis pollinated by the fig wasp,Blastophaga psenes.
Further information:Entomophily,Ornithophily,andReproductive coevolution in Ficus

Flowering plantsand the animals thatpollinatethem have co-evolved. Many plants that are pollinated byinsects(inentomophily),bats,orbirds(inornithophily) 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. Adaptivespeciationquickly gave rise to many diverse groups of plants, and, at the same time, corresponding speciation occurred in certaininsect 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,[72]where the plant species can only be pollinated by one species of insect.[73]

Pseudomyrmexant on bull thorn acacia (Vachellia cornigera) with Beltian bodies that provide the ants with protein[74]

Acacia ants and acacias

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Main article:Pseudomyrmex ferruginea

Theacacia ant(Pseudomyrmex ferruginea) is an obligate plant ant that protects at least five species of "Acacia" (Vachellia)[a]from preying insects and from other plants competing for sunlight, and the tree provides nourishment and shelter for the ant and its larvae.[74][75]

Seed dispersal

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Main article:Seed dispersal syndrome

Seed dispersal is the movement, spread or transport ofseedsaway from the parent plant. Plants have limited mobility and rely upon a variety ofdispersal vectorsto transport their propagules, including bothabioticvectors such as the wind and living (biotic) vectors like birds. In order to attract animals, these plants evolved a set of morphological characters such asfruitcolour, mass, and persistence correlated to particular seed dispersal agents.[76]For example, plants may evolve conspicuous fruit colours to attract avian frugivores, and birds may learn to associate such colours with a food resource.[77]

Rhizobia

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Main article:Rhizobia

Lichens

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Main article:Lichen

See also

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Notes

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  1. ^The acacia ant protects at least 5 species of "Acacia", now all renamed toVachellia:V. chiapensis,V. collinsii,V. cornigera,V. hindsiiandV. sphaerocephala.

References

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