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Many animal species cooperate with each other in mutual symbiosis. One example is the Ocellaris clownfish, which dwells among the tentacles of Ritteri sea anemones. The anemones provide the clownfish with protection from their predators (which cannot tolerate the stings of the anemone's tentacles), while the fish defend the anemones against butterflyfish (which eat anemones)

Cooperation (sometimes written as co-operation or coöperation[1]) is the process of groups of competition for selfish benefit. Many animal and plant species cooperate both with other members of their own species and with members of other species (symbiosis or mutualism).[2]


  • In humans 1
  • In animals 2
    • Kin selection 2.1
  • Cooperative systems 3
  • The Prisoner's Dilemma, the Ultimatum Game, and the Dictator Game. 4
  • See also 5
  • Notes 6
  • References 7
  • External links 8

In humans

Language allows humans to cooperate on a very large scale. Certain studies have shown that fairness affects human cooperation; individuals are willing to punish at their own cost (altruistic punishment) if they believe that they are being treated unfairly.[3][4] Sanfey, et al. conducted an experiment where 19 individuals were scanned using MRI while playing an Ultimatum Game in the role of the responder.[4] They were receiving offers from other human partners and from a computer partner. Remarkably, responders refused unfair offers from human partners at a significantly higher rate than those by a computer partner. The experiment also showed that altruistic punishment is associated with negative emotions that are being generated in unfair situations by the anterior insula of the brain.[4]

It has been observed that image scoring promotes cooperative behavior in situations where direct reciprocity is unlikely.[5] In situations where reputation and status are involved, humans tend to cooperate more.

In animals

Cooperation exists not only in humans but in animals as well. This behavior appears, however, to occur mostly between relatives. Spending time and resources assisting a related individual may at first seem destructive to the organism’s chances of survival but is actually beneficial over the long-term. Since relatives share part of their genetic make-up, enhancing each other’s chances of survival may actually increase the likelihood that the helper’s genetic traits will be passed on to future generations.[6]

Some researchers assert that cooperation is more complex than this. They maintain that helpers may receive more direct, and less indirect, gains from assisting others than is commonly reported. 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.[7]

Kin selection

One specific form of cooperation in animals is kin selection, which can be defined as animals helping to rear a relative’s offspring in order to enhance their own fitness.[7]

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. These two hypotheses both appear to be valid, at least in cichlid fish.[8]

Studies conducted on red wolves support previous researchers' [7] contention that helpers obtain both immediate and long-term gains from cooperative breeding. Researchers evaluated the consequences of red wolves’ decisions to stay with their packs for extended periods of time after birth. It was found that this "delayed dispersal," while it involved helping other wolves rear their offspring, extended male wolves’ life spans. These findings suggest that kin selection may not only benefit an individual in the long-term in terms of increased fitness but in the short-term as well through enhanced chance of survival [9]

Some research even suggests that certain species provide more help to the individuals with which they are more closely related. This phenomenon is known as kin discrimination.[10] In their meta-analysis, researchers compiled data on kin selection as mediated by genetic relatedness in 18 species, including the Western bluebird, Pied kingfisher, Australian magpie, and Dwarf Mongoose. They found that different species exhibited varying degrees of kin discrimination, with the largest frequencies occurring among those who have the most to gain from cooperative interactions.Griffin

Cooperative systems

Cooperation is a process by which the components of a system work together to achieve the global properties. In other words, individual components that appear to be "selfish" and independent work together to create a highly complex, greater-than-the-sum-of-its-parts system. Examples:

      • The components in a cell work together to keep it living.
    • *Neurons create thought and consciousness, other cells work together and communicate to produce multicellular organisms.
    • *Organisms form food chains and ecosystems.
    • *People form families, tribes, cities and nations.
    • *Atoms cooperate in a simple way, by combining to make up molecules.

Understanding the mechanisms that create cooperating agents in a system is one of the most important and least well understood phenomena in nature, though there has not been a lack of effort.

Individual action on behalf of a larger system may be coerced (forced), voluntary (freely chosen), or even unintentional, and consequently individuals and groups might act in concert even though they have almost nothing in common as regards interests or goals. Examples of that can be found in market trade, military wars, families, workplaces, schools and prisons, and more generally any institution or organization of which individuals are part (out of own choice, by law, or forced).

The Prisoner's Dilemma, the Ultimatum Game, and the Dictator Game.

For many years, the prisoner's dilemma game pointed out that even if all members of a group would benefit if all cooperate, 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. Results from experimental economics show that humans often act more cooperatively than strict self-interest would seem to dictate. While economic experiments require subjects to make relatively abstract decisions for small stakes, evidence from natural experiments for high stakes support the claim that humans act more cooperatively than strict self-interest would dictate.[11]

One reason may be that if the prisoner's dilemma situation is repeated (the iterated prisoner's dilemma), it allows non-cooperation to be punished more, and cooperation to be rewarded more, than the single-shot version of the problem would suggest. It has been suggested that this is one reason for the evolution of complex emotions in higher life forms,[12][13] who, at least as infants, and usually thereafter, cannot survive without cooperating – although with maturation they gain much more choice about the kinds of cooperation they wish to have.

In any case, two more recent games, the Ultimatum Game and the Dictator Game, showed that different context-related experimentations may lead to completely different results. In particular this is shown in the dictator's game, in which there are two human agents, one of which, "the ruler", is asked to decide the amount of wealth -usually in money-, between 100% and 0%, he/she is going to keep for him/herself and the one he/she is going to leave to the other human agent, which, in this game, is completely passive. Even if a "rational" economic behaviour would lead to keep the whole amount of good, it hasn't happened in experimentations, where human agents tend to share a variable amount of their goods anyway. These results were also considered a strong evidence of the absence of supposed "rational actors" in economy, especially from a neuroeconomics perspective.

See also


  1. ^
  2. ^
  3. ^
  4. ^ a b c
  5. ^
  6. ^ Hamilton, W.D. (1964). The Genetical Evolution of Social Behaviour I. Journal of Theoretical Biology, 7, 1-16.
  7. ^ a b c Clutton-Brock, T. (2002). Breeding together: Kin selection and mutualism in cooperative vertebrates. Science, 296(5565), 69-72. doi:10.1126/science.296.5565.69
  8. ^ Balshine-Earn, S., Neat, F.C., Reid, H., & Taborsky, M. (1998). Paying to stay or paying to breed? Field evidence for direct benefits of helping behavior in a cooperatively breeding fish. Behavioral Ecology, 9 (5), 432-438.
  9. ^ Sparkman, A. M., Adams, J. R., Steury, T. D., Waits, L. P., & Murray, D. L. (2011). Direct fitness benefits of delayed dispersal in the cooperatively breeding red wolf (Canis rufus). Behavioral Ecology, 22(1), 199-205. doi:10.1093/beheco/arq194
  10. ^ Griffin, A. S., & West, S. A. (2003). Kin Discrimination and the Benefit of Helping in Cooperatively Breeding Vertebrates. Science, 302(5645), 634-636. doi:10.1126/science.1089402
  11. ^
  12. ^
  13. ^


  • Herbert Gintis, Samuel Bowles, A Cooperative Species: Human Reciprocity and Its Evolution, Princeton University Press, 2011, ISBN 0-691-15125-3 (Reviewed in The Montreal Review)
  • Tom R. Tyler, "Why People Cooperate: The Role of Social Motivations", Princeton University Press, 2011, ISBN 9781400836666
  • Michael Tomasello, (2009), Why We Cooperate. MIT Press. ISBN 978-0-262-01359-8 (Reviewed in The Montreal Review)
  • Raimo Tuomela, "Cooperation, a philosophic study", Philosophical Studies Series, Vol. 82 Kluwer Academic Publishers, 2000, ISBN 978-94-015-9594-0
  • Robert Axelrod, The Complexity of Cooperation, Princeton Paperbacks, ISBN 0-691-01567-8
  • Robert Axelrod, The Evolution of Cooperation, Basic Books, ISBN 0-465-02121-2
  • Richard Dawkins (1990), The Selfish Gene, second edition – includes two chapters about the evolution of cooperation, ISBN 0-19-286092-5
  • Herbert Gintis, Samuel Bowles, Robert T. Boyd, Ernst Fehr (eds.), Moral Sentiments and Material Interests: The Foundations of Cooperation in Economic Life (Economic Learning and Social Evolution). MIT 2005
  • John McMurtry, "How Competition Goes Wrong." Journal of Applied Philosophy, 8(2): 200–210, 1991.
  • Dennis Rivers,, The Seven Challenges: A Workbook and Reader About Communicating More Cooperatively, fourth edition, 2005 – treats cooperation as a set of skills that can be improved.
  • M.J. van den Assem, D. van Dolder and R.H. Thaler (2010). "Split or Steal? Cooperative Behavior When the Stakes are Large"

External links

  • An Operation of Cooperation, A book about cooperation and the benefits of this path, as opposed to working alone.
  •, The Cooperation Project: Objectives, Accomplishments, and Proposals. Howard Rheingold's project with Institute for the Future.
  •, Cooperation platform for transport research (scientific)
  •, The Far Games, a list of games using theatrical improvisation to encourage collaboration and distributed leadership
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