Symbiosis

Symbiosis (Ancient Greek συμβίωσις symbíōsis: living with, companionship < σύν sýn: together; and βίωσις bíōsis: living)[2] is any type of a close and long-term biological interaction, between two organisms of different species. The two organisms, termed symbionts, can be either in a mutualistic, a commensalistic, or a parasitic relationship.[3] In 1879, Heinrich Anton de Bary defined symbiosis as "the living together of unlike organisms".
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 be obligatory, which means that one, or both of the organisms depend on each other for survival, or facultative (optional), when they can also subsist independently.
Symbiosis is also classified by physical attachment. Symbionts forming a single body live in conjunctive symbiosis, while all other arrangements are called disjunctive symbiosis.[4] When 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.[5][6]
Definition

The definition of symbiosis was a matter of debate for 130 years.[7] In 1877, Albert Bernhard Frank used the term symbiosis to describe the mutualistic relationship in lichens.[8][9] In 1878, the German mycologist Heinrich 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 persistent mutualisms, while others thought it should apply to all persistent biological interactions (in other words, to mutualism, commensalism, and parasitism, but excluding brief interactions such as predation). 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]
Types
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
Ectosymbiosis versus endosymbiosis
Contrastingly,
In endosymbiosis, the host cell lacks some of the nutrients which the
As the endosymbiont adapts to the host's lifestyle, the endosymbiont changes dramatically. There is a drastic reduction in its
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.[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 as food, water, and territory) used by both usually facilitates this type of interaction, although the competition can also be for other resources.[34]
Amensalism
Amensalism is a non-symbiotic, asymmetric interaction where one species is harmed or killed by the other, and one is unaffected by the other.
The term amensalism is often used to describe strongly asymmetrical competitive interactions, such as between the
Commensalism
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 word
Commensal relationships may involve one organism using another for transportation (
Mutualism

Mutualism or interspecies reciprocal altruism is a long-term relationship between individuals of different species 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.
Many
An example of mutualism is the relationship between the
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, and both quickly retreat into the burrow.[45] Different species of gobies (Elacatinus spp.) also clean up ectoparasites in other fish, possibly another kind of mutualism.[46]
A spectacular example of obligate mutualism is the relationship between the
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 encrusting
Parasitism
In a parasitic relationship, the parasite benefits while the host is harmed.
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.[52][53] In contrast, Müllerian mimicry is mutually beneficial as all participants are both models and mimics.[54][55] 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. [56]
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).[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 of cleaner fish – notably wrasses, 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]
Role in evolution

Symbiosis is increasingly recognized as an important selective force behind evolution;
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]
Hologenome development and evolution
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]
Symbiogenesis
One hypothesis for the origin of the nucleus in
The biologist
Major examples of co-evolutionary relationships
Mycorrhiza
About 80% of

Pollination

Acacia ants and acacias
The
Seed dispersal
Seed dispersal is the movement, spread or transport of
Rhizobia
Lichens
See also
Notes
- ^ 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.
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External links
Media related to Symbiosis at Wikimedia Commons
The dictionary definition of symbiosis at Wiktionary
- TED-Education video – Symbiosis: a surprising tale of species cooperation