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ANTH1014; ZFZY0; TA Name: Kathleen Bryson; Word Count: 1500

Why do selfish organisms cooperate?

A study conducted in 2011 by a team of scientists on 12 elephants at the Thai Elephant Conservation Center in Lampang, Thailand, revealed that elephants are highly cooperative. Working in pairs, in order for the animals to get a bucket of corn, they had to pull two ends of the same rope at the same time, so that a table with the corn slid within their reach. If one end of rope was pulled before or without the other, the whole rope slipped, leaving the elephants with nothing. Interestingly, they quickly learned to cooperate and none of them tried to cheat, that is, to try and reach for the corn on its own (Choi 2011). In intelligent mammals such as elephants, cooperation involves complex cognition tasks. But cooperation can also be found in species varying from bees to primates. Why should it have evolved in the first place, then? At first glance, a simplistic explanation such as Cooperation means mutual help, therefore benefits for both sides, so why not? may seem logical. Yet it ignores both the fact that there are also costs to helping another individual, and the fact that natural selection is unlikely to favour altruist genes. Therefore, cooperation must somehow aid selfishness (Boyd&Silk 2006). I argue that the idea of altruism per se has to be replaced with that of pseudo-altruism, which is ultimately selfish. I first draw on a few intriguing anomalies such as altruistic suicide in bees or the occasions on which victorious deer do not kill their rivals, just to show how groupselectionist answers deal with them; then, I explain the most widely accepted theory about the mechanism that lies behind altruism, which is kin selection; after that, I summarize the points made by an opponent of kin selection; another aspect of pseudo-altruism as an evolutionary trait

is how it can explain why humans evolved as social and moral beings so I look at moralistic aggression, gratitude, friendship etc.; in the end, I conclude that regardless of what mechanism is at the core of altruism, it is not favoured by natural selection, and therefore it is rare and maladaptive. Altruistic behaviour is a Darwinian paradox, as it benefits others at a personal cost to the individual. Mutualism is when this behaviour benefits the actor as well as the recipient. It is fragile, considering that slacking can be beneficial for the individuals, as found in coalitions amongst male baboons. Odd behaviours such as when bees are willing to die to defend their hive, or when deer who win a fight do not kill their rivals (Lorenz 1963), have been explained through group-selection, which states that the individual behaviour aids the preservation of a group, population or species (Wynne-Edwards 1962). Nevertheless, natural selection depends on genetic variability, which is greater within a group than between groups. Genes promoting selfishness would spread much faster than those promoting altruism. Then what is really going on? In order for natural selection to favour altruism, the benefits to the recipient have to outweigh the costs suffered by the actor. How could that be? If altruism is performed among individuals more likely to carry a gene for altruism, then that gene will increase in frequency. The probability of individuals sharing a gene for altruism is equivalent to asking what the average degree of relatedness between two individuals is. So an individual is 100% related to himself, 50% related to his parents and full-siblings, 25% related to his grandparents and half-siblings, 12.5% to his great-grandparents or cousins etc. Mathematically, it is easy to see how the average degree of relatedness multiplied by the benefits to the recipient have to outweigh the costs in order for an altruistic behaviour to result in increased fitness. This is why altruism is not expected to be found amongst non-related

individuals (Hamilton 1964). Back to a group-selectionist view, groups containing altruistic individuals always out-compete those with no or fewer such individuals. But within a group, altruistic individuals are disadvantaged. Following this logic, it is reasonable that recognition mechanisms should have evolved so as to, on the one hand, enable individuals to distinguish their relatives from non-related individuals, on the other hand to enable them to maintain helpful social relationships. West et al. (2007) distinguish between genetic cues (smell) and environmental ones (shared environment or previous association), stating that the latter are more effective than the former, which would most likely reduce precisely the variation required for kin selection to occur. Another possible mechanism would be the green-beard effect (a term coined by Richard Dawkins in 1976), which states that a trait has to be perceptible, has to enable recognition of the same trait in others and a preferential treatment to those recognized. On the other hand, the importance of social relationships stems from how they aid cooperation. For instance, grooming cultivates social bonds between individuals useful to one another for coalitions, such as in male red howler monkeys. They live in groups consisting of two to four females and one or two males, which really makes mating a hard task. Males work together to gain access to breeding females by evicting male residents, and by mate guarding. According to Trivers (1985), kin selection explains parents-offspring rivalry: selection favours mothers who provide less than their infants desire, and infants that demand more than their mothers are willing to give. Not everybody agrees. Goldschmidt (2005), focusing on the mother-child relationship and cooperation rather than on couples or social groups, developed the theory of affect hunger (the craving for attention and affection as means of manipulating other

people to gain access to various resources) as a neuro-anthropological alternative to kin selection. He has no problem with arguing that people are selfish, but only with arguing that altruism is the right way to be selfish. He challenges kin selection stating that there is hardly any consideration of neural mechanisms involved in the process it all sums up to the assumption that the gene causes altruistic behaviour if it is favourable to its transmission. Moreover, altruism should be even more selective after all, human beings often behave altruistically towards nonrelated individuals! Acknowledging the kin-selectionist theory which explains indiscriminate altruism through the fact that our ancestors were generally in the exclusive presence of their kin, he does not find this explanation reasonable for altruism extended to other species. Another interesting point he makes is that kin selection takes for granted the assumption that organic variation easily produces adaptations suitable to any environment, and that all traits are necessarily adaptive, when in fact selection acts upon a whole organism and given that traits are often genetically connected, selection might act on another trait produced by the same gene. Despite the fact that the two views mentioned above clash when it comes to explaining how cooperation evolved, both of them acknowledge the inexistence of pure altruism as an adaptive behavioural trait. I would argue that when we rush to save somebody in urgent need, we act instinctively without considering any beneficial outcome to our inclusive fitness. Even so, this proves the existence of pure altruism, but not that it would occur on a daily basis or be favoured by natural selection. To outline wider socio-psychological implications of our adaptations towards cooperation, I now turn to an article by Trivers (1985) which explains the evolution of sociality and morality as beneficial for cooperation. He states that our tendency to form relationships and

to act altruistically towards friends and towards the people we like goes both ways: the more we like them, the more altruistically we behave towards them and the more altruistic they are, the more we like them. I believe that our social complexities confirm this view: a lot of people would rather help their very close friends rather than an estranged relative. Regarding moralistic aggression, always preceded by a sense of fairness, it is yet another mechanism ensuring the strength and durability of social relationships, and the punishment and ostracizing of cheaters. Even gratitude, sympathy, guilt and reparative altruism have evolved in order to preserve and repair social bonds. This is especially important in the human species, where systems of multiparty altruism may operate simultaneously, so that an individual does not gain benefits from the individual aided, but from a third-party individual. I have argued that cooperation, especially altruism, is an apparent Darwinian paradox, for it costs the actor who could be better off not cooperating at all. Group selection is not a viable explanation, as it would destroy the very variability that laid its foundation. While kin selection is most widely acknowledged, complementary views still need to be considered, such as Goldschmidts affect hunger theory. As a concluding remark, I state that pure altruism is not favoured by natural selection and, in this sense, can be pictured as a maladaptive behaviour. However, pseudo-altruism and other types of cooperation are favoured, and understanding their further influences on our sociality and morality may help us reassess just how good and selfless we think we are.

References
BOYD, R. AND SILK, J. B. 2006. How Humans Evolved. University of California, Los Angeles. CHOI, C. Q. 2011. Elephants cooperate, proving how smart they really are. http://www.livescience.com/13108-elephants-cooperate-intelligent-behavior.html DAWKINS, R. 1976. The Selfish Gene. Oxford University Press. GOLDSCHMIDT, W. 2005. The Bridge to Humanity. How Affect Hunger Trumps the Selfish Gene. Oxford University Press. HAMILTON, W. D. 1996. The Evolution of Social Behaviour. Journal of Theoretical Biology. LORENZ, K. 1963. On aggression. Harcourt and Brace Company. TRIVERS, R. 1985. Social Evolution. Menlo Park: Benjamin Cummings. WEST, S. A.; GRIFFIN, A. S.; GARDNER, A. 2007. Evolutionary explanations for cooperation. Current Biology 17: R661-R672. WYNNE-EDWARDS, V. 1962. Animal Dispersion in Relation to Social Behaviour. Edinburgh: Oliver & Boyd.

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