Cooperation (evolution)
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In evolution, cooperation is the process where groups of organisms work or act together for common or mutual benefits. It is commonly defined as any adaptation that has evolved, at least in part, to increase the reproductive success of the actor's social partners.[1] For example, territorial choruses by male lions discourage intruders and are likely to benefit all contributors.[2]
This process contrasts with intragroup competition where individuals work against each other for selfish reasons. Cooperation exists not only in humans but in other animals as well. The diversity of taxa that exhibits cooperation is quite large, ranging from
In animals
Cooperation in animals appears to occur mostly for direct benefit or between relatives. Spending time and resources assisting a related individual may at first seem destructive to an organism's chances of survival but is actually beneficial over the long-term. Since relatives share part of the helper's genetic make-up, enhancing each individual's chance of survival may actually increase the likelihood that the helper's genetic traits will be passed on to future generations.[3]
However, some researchers, such as ecology professor Tim Clutton-Brock, assert that cooperation is a more complex process. They state that helpers may receive more direct, and less indirect, gains from assisting others than is commonly reported. These gains include protection from predation and increased reproductive fitness. Furthermore, they insist that cooperation may not solely be an interaction between two individuals but may be part of the broader goal of unifying populations.[4]
Prominent biologists, such as Charles Darwin, E. O. Wilson, and W. D. Hamilton, have found the evolution of cooperation fascinating because natural selection favors those who achieve the greatest reproductive success while cooperative behavior often decreases the reproductive success of the actor (the individual performing the cooperative behavior). Hence, cooperation seemed to pose a challenging problem to the theory of natural selection, which rests on the assumption that individuals compete to survive and maximize their reproductive successes.[2] Additionally, some species have been found to perform cooperative behaviors that may at first sight seem detrimental to their own evolutionary fitness. For example, when a ground squirrel sounds an alarm call to warn other group members of a nearby coyote, it draws attention to itself and increases its own odds of being eaten.[5] There have been multiple hypotheses for the evolution of cooperation, all of which are rooted in Hamilton's models based on inclusive fitness. These models hypothesize that cooperation is favored by natural selection due to either direct fitness benefits (mutually beneficial cooperation) or indirect fitness benefits (altruistic cooperation).[6] As explained below, direct benefits encompass by-product benefits and enforced reciprocity, while indirect benefits (kin selection) encompass limited dispersal, kin discrimination and the greenbeard effect.
Kin selection
One specific form of cooperation in animals is kin selection, which involves animals promoting the reproductive success of their kin, thereby promoting their own fitness.[4][nb 1]
Different theories explaining kin selection have been proposed, including the "pay-to-stay" and "territory inheritance" hypotheses. The "pay-to-stay" theory suggests that individuals help others rear offspring in order to return the favor of the breeders allowing them to live on their land. The "territory inheritance" theory contends that individuals help in order to have improved access to breeding areas once the breeders depart.[9]
Studies conducted on red wolves support previous researchers' contention that helpers obtain both immediate and long-term gains from cooperative breeding.[4] Researchers evaluated the consequences of red wolves' decisions to stay with their packs for extended periods of time after birth. While delayed dispersal helped other wolves' offspring, studies also found that it extended male helper wolves' life spans. This suggests that kin selection may not only benefit an individual in the long-term through increased fitness but also in the short-term through increased survival chances.[10]
Some research suggests that individuals provide more help to closer relatives. This phenomenon is known as kin discrimination.
In plants
Cooperation exists not only in animals but also in plants. In a greenhouse experiment with Ipomoea hederacea, a climbing plant, results show that kin groups have higher efficiency rates in growth than non-kin groups do. This is expected to rise out of reduced competition within the kin groups.[12]
Explanation
The inclusive fitness theory provides a good overview of possible solutions to the fundamental problem of cooperation. The theory is based on the hypothesis that cooperation helps in transmitting underlying genes to future generations either through increasing the reproductive successes of the individual (direct fitness) or of other individuals who carry the same genes (indirect fitness). Direct benefits can result from simple by-product of cooperation or enforcement mechanisms, while indirect benefits can result from cooperation with genetically similar individuals.[3]
Direct fitness benefits
This is also called mutually beneficial cooperation as both actor and recipient depend on direct fitness benefits, which are broken down into two different types:
By-product benefit arises as a consequence of social partners having a shared interest in cooperation. For example, in meerkats, larger group size provides a benefit to all the members of that group by increasing survival rates, foraging success and conflict wins.
Prisoner's Delight, another term to describe by-product benefit, is a term coined by
It has been shown that
Cooperation is maintained in situations where
Enforcement can also be mutually beneficial, and is often called reciprocal cooperation because the act of cooperation is preferentially directed at individuals who have helped the actor in the past (directly), or helped those who have helped the actor in the past (indirectly).[18]
Indirect fitness benefits
The second class of explanations for cooperation is indirect fitness benefits, or
Hamilton originally suggested that high relatedness could arise in two ways: direct kin recognition between individuals or limited dispersal, or population viscosity, which can keep relatives together.[19] The easiest way to generate relatedness between social partners is limited dispersal, a mechanism in which genetic similarity correlates with spatial proximity. If individuals do not move far, then kin usually surrounds them. Hence, any act of altruism would be directed primarily towards kin. This mechanism has been shown in Pseudomonas aeruginosa bacteria, where cooperation is disfavored when populations are well mixed, but favored when there is high local relatedness.[20]
Kin discrimination also influences cooperation because the actor can give aid preferentially towards related partners. Since kin usually share common genes, it is thought that this nepotism can lead to genetic relatedness between the actor and the partner's offspring, which affects the cooperation an actor might give.
This mechanism is similar to what happens with the
Multi-level selection
Multi-level selection theory suggests that selection operates on more than one level: for example, it may operate at an atomic and molecular level in cells, at the level of cells in the body, and then again at the whole organism level, and the community level, and the species level. Any level which is not competitive with others of the same level will be eliminated, even if the level below is highly competitive. A classic example is that of genes which prevent cancer. Cancer cells divide uncontrollably, and at the cellular level, they are very successful, because they are (in the short term) reproducing very well and out competing other cells in the body. However, at the whole organism level, cancer is often fatal, and so may prevent reproduction. Therefore, changes to the genome which prevent cancer (for example, by causing damaged cells to act co-operatively by destroying themselves) are favoured. Multi-level selection theory contends that similar effects can occur, for example, to cause individuals to co-operate to avoid behaviours which favour themselves short-term, but destroy the community (and their descendants) long term.
Market effect
One theory suggesting a mechanism that could lead to the evolution of co-operation is the "market effect" as suggested by Noe and Hammerstein.
This mechanism can be relied to both within a species or social group and within species systems. It can also be applied to a multi-partner system, in which the owner of a resource has the power to choose its co-operation partner. This model can be applied in natural systems (examples exist in the world of apes, cleaner fish, and more). Easy for exemplifying, though, are systems from international trading. Arabic countries control vast amounts of oil, but seek technologies from western countries. These in turn are in need of Arab oil. The solution is co-operation by trade.
Symbiosis
Symbiosis refers to two or more biological species that interact closely, often over a long period of time. Symbiosis includes three types of interactions—mutualism, commensalism, and parasitism—of which only mutualism can sometimes qualify as cooperation. Mutualism involves a close, mutually beneficial interaction between two different biological species, whereas "cooperation" is a more general term that can involve looser interactions and can be interspecific (between species) or intraspecific (within a species). In commensalism, one of the two participating species benefits, while the other is neither harmed nor benefitted. In parasitism, one of the two participating species benefits at the expense of the other.
Symbiosis may be obligate or facultative. In obligate symbiosis, one or both species depends on the other for survival. In facultative symbiosis, the symbiotic interaction is not necessary for the survival of either species.
Two special types of symbiosis include endosymbiosis, in which one species lives inside of another, and ectosymbiosis, in which one species lives on another.
Mutualism
Mutualism is a form of symbiosis in which both participating species benefit.
A classic example of mutualism is the interaction between rhizobia soil bacteria and legumes (Fabaceae). In this interaction, rhizobia bacteria induce root nodule formation in legume plants via an exchange of molecular signals.[23] Within the root nodules, rhizobia fix atmospheric nitrogen into ammonia using the nitrogenase enzyme. The legume benefits from a new supply of usable nitrogen from the rhizobia, and the rhizobia benefits from organic acid energy sources from the plant as well as the protection provided by the root nodule. Since the rhizobia live within the legume, this is an example of endosymbiosis, and since both the bacteria and the plant can survive independently, it is also an example of facultative symbiosis.
Not all examples of mutualism are also examples of cooperation. Specifically, in by-product mutualism, both participants benefit, but cooperation is not involved. For example, when an elephant defecates, this is beneficial to the elephant as a way to empty waste, and it is also beneficial to a dung beetle that uses the elephant's dung. However, neither participant's behavior yields a benefit from the other, and thus cooperation is not taking place.[25]
Hidden benefits
Hidden benefits are benefits from cooperation that are not obvious because they are obscure or delayed. (For example, a hidden benefit would not involve an increase in the number of offspring or offspring viability.)
One example of a hidden benefit involves Malarus cyaneus, the
Another example of a hidden benefit is indirect reciprocity, in which a donor individual helps a beneficiary to increase the probability that observers will invest in the donor in the future, even when the donor will have no further interaction with the beneficiary.
In a study of 79 students, participants played a game in which they could repeatedly give money to others and receive from others. They were told that they would never interact with the same person in the reciprocal role. A player's history of donating was displayed at each anonymous interaction, and donations were significantly more frequent to receivers who had been generous to others in earlier interactions.[26] Indirect reciprocity has only been shown to occur in humans.[27]
Prisoner's dilemma
Even if all members of a group benefit from cooperation, individual self-interest may not favor cooperation. The prisoner's dilemma codifies this problem and has been the subject of much research, both theoretical and experimental. In its original form the prisoner's dilemma game (PDG) described two awaiting trial prisoners, A and B, each faced with the choice of betraying the other or remaining silent. The "game" has four possible outcomes: (a) they both betray each other, and are both sentenced to two years in prison; (b) A betrays B, which sets A free and B is sentenced to four years in prison; (c) B betrays A, with the same result as (b) except that it is B who is set free and the other spends four years in jail; (d) both remain silent, resulting in a six-month sentence each. Clearly (d) ("cooperation") is the best mutual strategy, but from the point of view of the individual betrayal is unbeatable (resulting in being set free, or getting only a two-year sentence). Remaining silent results in a four-year or six-month sentence. This is exemplified by a further example of the PDG: two strangers attend a restaurant together and decide to split the bill. The mutually best ploy would be for both parties to order the cheapest items on the menu (mutual cooperation). But if one member of the party exploits the situation by ordering the most expensive items, then it is best for the other member to do likewise. In fact, if the fellow diner's personality is completely unknown, and the two diners are unlikely ever to meet again, it is always in one's own best interests to eat as expensively as possible. Situations in nature that are subject to the same dynamics (rewards and penalties) as the PDG define cooperative behavior: it is never in the individual's fitness interests to cooperate, even though mutual cooperation rewards the two contestants (together) more highly than any other strategy.[28] As described in the Nash equilibrium, cooperation cannot evolve under these circumstances.
However, in 1981 Axelrod and Hamilton[29] noted that if the same contestants in the PDG meet repeatedly (in the so-called iterated prisoner's dilemma game, IPD) then tit-for-tat (foreshadowed by Robert Trivers' 1971 reciprocal altruism theory[30]) is a robust strategy which promotes altruism.[28][29][31] In "tit-for-tat" both players' opening moves are cooperation. Thereafter each contestant repeats the other player's last move, resulting in a seemingly endless sequence of mutually cooperative moves. However, mistakes severely undermine tit-for-tat's effectiveness, giving rise to prolonged sequences of betrayal, which can only be rectified by another mistake. Since these initial discoveries, all the other possible IPD game strategies have been identified (16 possibilities in all, including, for instance, "generous tit-for-tat", which behaves like "tit-for-tat", except that it cooperates with a small probability when the opponent's last move was "betray".[32]), but all can be outperformed by at least one of the other strategies, should one of the players switch to such a strategy. The result is that none is evolutionarily stable, and any prolonged series of the iterated prisoner's dilemma game, in which alternative strategies arise at random, gives rise to a chaotic sequence of strategy changes that never ends.[28][33][34]
Results from experimental economics show, however, that humans often act more cooperatively than strict self-interest would dictate.[35]
Evolutionary mechanisms suggesting that reciprocity is the result, not the cause, of the evolution of cooperation
In the light of the iterated prisoner's dilemma game and the reciprocal altruism theory failing to provide full answers to the evolutionary stability of cooperation, several alternative explanations have been proposed.
There are striking parallels between cooperative behavior and exaggerated
There is an alternate strategy for identifying fit mates which does not rely on one gender having exaggerated sexual ornaments or other handicaps, but is probably generally applicable to most, if not all sexual creatures. It derives from the concept that the change in appearance and functionality caused by a
History of cooperation research
One of the first references to animal cooperation was made by Charles Darwin, who noted it as a potential problem for his theory of natural selection.[48] In most of the 19th century, intellectuals like Thomas Henry Huxley and Peter Kropotkin debated fervently on whether animals cooperate with one another and whether animals displayed altruistic behaviors.[49]
In the late 1900s, some early research in animal cooperation focused on the benefits of group-living. While living in a group produces costs in the form of increased frequency of predator attacks and greater mating competition, some animals find that the benefits outweigh the costs. Animals that practice group-living often benefit from assistance in parasite removal, access to more mates, and conservation of energy in foraging.
In the past, simple game theory models, such as the classic cooperative hunting and Prisoner's dilemma models, were used to determine decisions made by animals in cooperative relationships. However, complicated interactions between animals have required the use of more complex economic models such as the Nash equilibrium. The Nash equilibrium is a type of non-cooperative game theory that assumes an individual's decision is influenced by its knowledge of the strategies of other individuals. This theory was novel because it took into consideration the higher cognitive capabilities of animals.[51][52] The evolutionarily stable strategy is a refined version of the Nash equilibrium in that it assumes strategies are heritable and are subject to natural selection. Economic models are useful for analyzing cooperative relationships because they provide predictions on how individuals act when cooperation is an option. Economic models are not perfect, but they provide a general idea of how cooperative relationships work.
Contrary to the mainstream dogma, a recently published article.[53] using agent-based models demonstrates that several crucial mechanisms, such as kin selection, punishment, multilevel selection, and spatial structure, cannot rescue the evolution of cooperation. The new findings revive a long-standing puzzle in the evolution theory. In addition, the work has potential therapeutic benefits for numerous incurable diseases.
See also
- Agreeableness
- Collaboration
- Dunbar's number
- Evolution of cooperation
- Management cybernetics
- Microbial Cooperation
- Mutual Aid: A Factor of Evolution by Peter Kropotkin
- Polytely
- Teamwork
Notes
- ^ An individual's gene complement (or genome) can be represented by the letters of the alphabet. Each letter is represented twice: A1 and A2. This individual's genome therefore consists of 52 genes. The subscript indicates from which parent that copy of A has come. Mostly the two copies are identical, but occasionally they differ slightly. When this individual reproduces sexually, one or other copy of A (chosen randomly) is passed on to offspring-1, who gets its other copy of A from the sexual partner. The same happens with genes B, C, D, ..., Z. If we denote the two sexual partners by means of subscripts "m" and "f", then the genome of the offspring they produce might consist of Am2/Af1, Bm2/Bf2, Cm1/Cf1, Dm1/Df1 ... Zm1/Zf2. Each parent has contributed exactly half of the offspring's genome. So individual "m" shares only half of its genome with its offspring. Suppose individuals "m" and "f" produce a second offspring (offspring-2), whose genome is determined in exactly the same manner. There is a coin-flip 50% probability that offspring-2 will inherit the same copy of A from "m" as offspring-1 did (i.e. Am2). This also applies to gene B and so on through the alphabet. If a coin-flip "heads" means that gene X is the same in offspring-1 as it is in offspring-2, then in 26 flips of the coin approximately half are going to be "heads" and the rest "tails", i.e. half the genes inherited from parent "m" will be the same in the two offspring. The same will happen to the genes inherited from parent "f". Thus of the 52 genes inherited from the two parents, on average, 13 + 13 = 26 (or half) will be identical in the two sibs. Thus sibs are genetically as similar to one another as a parent is to an offspring.[7][8] From a evolutionary genetic point of view it is therefore as advantageous to help with the upbringing of full sibs as it is to produce and raise one's own offspring.
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