Symbiosis

Symbiosis (from Greek συμβίωσις, symbíōsis, "living with, companionship, camaraderie", from σύν, sýn, "together", and βίωσις, bíōsis, "living")^[2] is any type of a close and long-term biological interaction between two biological organisms of different species, termed symbionts, be it mutualistic, commensalistic, or parasitic.^[3] In 1879, Heinrich Anton de Bary defined it as "the living together of unlike organisms". The term is sometimes used in the more restricted sense of a mutually beneficial interaction in which both symbionts contribute to each other's support.^[3]

Symbiosis can be obligatory, which means that one or more of the symbionts depend on each other for survival, or facultative (optional), when they can generally live independently.

Symbiosis is also classified by physical attachment. When symbionts form a single body it is called 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]}

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

Competition can be defined as an interaction between organisms or species, in which the fitness of one is lowered by the presence of another.^[26] 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.^{[27][page needed]}

Mutualism

Mutualism or interspecies reciprocal altruism is a long-term relationship between individuals of different species where both individuals benefit.^[28] Mutualistic relationships may be either obligate for both species, obligate for one but facultative for the other, or facultative for both.

Many

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.^[33] Different species of gobies (Elacatinus spp.) also clean up ectoparasites in other fish, possibly another kind of mutualism.^[34]

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.[36]

A facultative symbiosis is seen in encrusting

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.^[47][48] In contrast, Müllerian mimicry is mutually beneficial as all participants are both models and mimics.^[49][50] 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. ^[51]

Amensalism

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).^[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 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.^[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.^[60]

Co-evolution and hologenome theory

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.^[64] The hologenome theory relates to the holobiont and symbionts genome together as a whole.^[65] Microbes live everywhere in and on every multicellular organism.^[66] 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.^[64]

Symbiogenesis

One hypothesis for the origin of the nucleus in

Pollination

Acacia ants and acacias

The

Seed dispersal is the movement, spread or transport of

Notes

References