In acleaning symbiosis, theclownfish feeds 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. This means that they each benefit each other in some way.[3]
Symbiosis can beobligatory, which means that one, or both of the organisms 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 inconjunctive symbiosis, while all other arrangements are calleddisjunctive symbiosis.[4] When one organism lives on the surface of another, such ashead lice on humans, it is calledectosymbiosis; when one partner lives inside the tissues of another, such asSymbiodinium withincoral, it is termedendosymbiosis.[5][6]
Diagram of the six possible types of symbiotic relationship, from mutual benefit to mutual harm.
The definition ofsymbiosis was a matter of debate for 130 years.[7] In 1877,Albert Bernhard Frank used the termsymbiosis to describe the mutualistic relationship inlichens.[8][9] In 1878, the GermanmycologistHeinrich Anton de Bary defined 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 Haskell proposed an integrative approach with a classification of "co-actions",[14] later adopted by biologists as "interactions".[15][16][17][18]
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 andphotosynthetic symbionts, 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]
In endosymbiosis, the host cell lacks some of the nutrients which theendosymbiont 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 viavertical transmission (heredity).[28]
As the endosymbiont adapts to the host's lifestyle, the endosymbiont changes dramatically. There is a drastic reduction in itsgenome size, as many genes are lost during the process ofmetabolism, andDNA repair and recombination, while important genes participating in the DNA-to-RNAtranscription, proteintranslation 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 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 typephenotype via a recombination process is calledMuller's ratchet phenomenon. Muller's ratchet phenomenon, together with less effective population sizes, leads to an accretion of deleteriousmutations in the non-essential genes of the intracellular bacteria.[29] This can be due to lack ofselection mechanisms prevailing in the relatively "rich" host environment.[30][31]
Competition can be defined as an interaction betweenorganisms or species, in which thefitness of one is lowered by the presence of another.[32] Competition can also occur between cells within the same organism which is why older cells are usually eliminated from tissues. This allows the organism to stay as healthy as possible by constantly eliminating old cells and making new ones.[33]Limited supply 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.[34]
Theblack walnut secretes 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.[35][36] 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 thesapling 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 isJuglans nigra (black walnut), secretingjuglone, a substance which destroys many herbaceous plants within its root zone.[37]
The termamensalism is often used to describe strongly asymmetrical competitive interactions, such as between theSpanish ibex andweevils of the genusTimarcha 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.[38]
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).[39][40]
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 crabs usinggastropod shells to protect their bodies, and spiders building their webs onplants.
Mutualism or interspeciesreciprocal altruism is a long-term relationship between individuals of differentspecies where both individuals benefit.[39] Mutualistic relationships may be either obligate for both species, obligate for one but facultative for the other, or facultative for both.
Manyherbivores have mutualisticgut flora to help them digest plant matter, which is more difficult to digest than animal prey.[5] This gut flora comprises cellulose-digestingprotozoans or bacteria living in the herbivores' intestines.[41]Coral reefs result from mutualism between coral organisms and various algae living inside them.[42] Most land plants and land ecosystems rely on mutualism between the plants, whichfix carbon from the air, andmycorrhyzal fungi, which help in extracting water and minerals from the ground.[43]
An example of mutualism is the relationship between theocellaris clownfish that dwell among thetentacles ofRitteri 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 specialmucus on the clownfish protects it from the stinging tentacles.[44]
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.[45] Different species of gobies (Elacatinus spp.) alsoclean up ectoparasites in other fish, possibly another kind of mutualism.[46]
A spectacular example of obligate mutualism is the relationship between thesiboglinidtube worms and symbioticbacteria that live athydrothermal vents andcold seeps. The worm has no digestive tract and is wholly reliant on its internal symbionts for nutrition. The bacteria oxidize eitherhydrogen 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 atdeep-sea hydrothermal vents and cold seeps in all of the world's oceans.[47]
Mutualism improves both organism's competitive ability and will outcompete organisms of the same species that lack the symbiont.[48]
A facultative symbiosis is seen in encrustingbryozoans andhermit 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.[49]
In a parasitic relationship, the parasite benefits while the host is harmed.[50] Parasitism takes many forms, fromendoparasites that live within the host's body toectoparasites andparasitic castrators that live on its surface andmicropredators like 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.[51]
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, todeceive a 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.[52][53] In contrast,Müllerian mimicry is mutually beneficial as all participants are both models and mimics.[54][55] For example, different species ofbumblebee mimic each other, with similar warning coloration in combinations of black, white, red, and yellow, and all of them benefit from the relationship.[56]
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).[57] 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.[58] In a supreme situation, the host species (fish or marine life) will display itself at a designated station deemed the "cleaning station".[59]
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.[60] 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.[60]
Symbiosis is increasingly recognized as an important selective force behind evolution;[5][61] many species have a long history of interdependentco-evolution.[62]
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.[63]
Mutualistic symbiosis can sometimes evolve fromparasitism orcommensalism, Fungi's relationship to plants in the form ofmycelium evolved from parasitism andcommensalism. Under certain conditions species of fungi previously in a state of mutualism can turn parasitic on weak or dying plants.[64] Likewise the symbiotic relationship ofclown fish andsea anemones emerged from a commensalist relationship.[65][66][67]
Evolution originated from changes in development where variations within species are selected for or against because of the symbionts involved.[68] The hologenome theory relates to the holobiont and symbionts genome together as a whole.[69] Microbes live everywhere in and on every multicellular organism.[70] 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.[68]
One hypothesis for the origin of the nucleus ineukaryotes (plants, animals, fungi, andprotists) is that it developed from asymbiogenesis between bacteria and archaea.[5][71][72] 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.[73] Evidence for this includes the fact thatmitochondria andchloroplasts divide independently of the cell, and that these organelles have their own genome.[74]
Flowering plants and the animals thatpollinate them 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. Adaptivespeciation quickly 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,[77] where the plant species can only be pollinated by one species of insect.[78]
Pseudomyrmex ant on bull thorn acacia (Vachellia cornigera) with Beltian bodies that provide the ants with protein[79]
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.[79][80]
Seed dispersal is the movement, spread or transport ofseeds away from the parent plant. Plants have limited mobility and rely upon a variety ofdispersal vectors to transport their propagules, including bothabiotic vectors such as the wind and living (biotic) vectors like birds. In order to attract animals, these plants evolved a set of morphological characters such asfruit colour, mass, and persistence correlated to particular seed dispersal agents.[81] For example, plants may evolve conspicuous fruit colours to attract avian frugivores, and birds may learn to associate such colours with a food resource.[82]
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