Evolutionary, neurobiological, gene-based solution of the ideological "puzzle" of human altruism and cooperation.

Despite hundreds of published articles about humankind's eusocial behaviours, most scholars still regard the origin of human altruism and cooperation as an enduring puzzle, because it seems incompatible with two central tenets of evolution, namely, the competition between individuals and the consequent selective advantages of selfish traits. This "puzzle", however, rather than being due to insurmountable scientific difficulties, is to be attributed to two powerful ideologies, which are politically opposite, but nevertheless concurred to prevent scholars from solving it. One ideology rejects the concept of genetic determinism, whereas the other dislikes the concept of group selection. As a consequence, these widespread ideologies, which are common in the scientific community, too, kept scholars from realising that the puzzle of human altruism and cooperation can only be solved by proposing a theoretical model that is based precisely on both genetic determinism and group selection. This model, which was never advanced in published papers, is presented here. This article also proposes to regard ancestral environments as determinants of human eusociality. By contrast, virtually all previous articles about it leave primitive habitats unmentioned. To support the hypothesis that human unselfish behaviours represent genetically conserved traits that evolved ancestrally, not products of cultural transmission, this paper also discusses six groups of arguments in the section "Genes versus culture". Finally, this article advances a purely genetic evolutionary explanation for the uniqueness of human eusociality, thereby challenging prevailing cultural explanations for the incomparably developed levels of cooperation in humankind, which are observed in no other social species.

[1]  Pavel Stopka,et al.  Sociology: The puzzle of human cooperation. , 2003, Nature.

[2]  Angel Sánchez,et al.  Altruism may arise from individual selection. , 2004, Journal of theoretical biology.

[3]  E. Wilson Genomics: How to make a social insect , 2006, Nature.

[4]  K. Hawkes,et al.  Hadza meat sharing. , 2001, Evolution and human behavior : official journal of the Human Behavior and Evolution Society.

[5]  R. Plomin The role of inheritance in behavior. , 1990, Science.

[6]  P. Taylor Altruism in viscous populations — an inclusive fitness model , 1992, Evolutionary Ecology.

[7]  R. James R. Blair,et al.  The neural basis of implicit moral attitude—An IAT study using event-related fMRI , 2006, NeuroImage.

[8]  H. Bandelt,et al.  Genetic evidence of an early exit of Homo sapiens sapiens from Africa through eastern Africa , 1999, Nature Genetics.

[9]  L. Keller,et al.  The evolution of cooperation and altruism – a general framework and a classification of models , 2006, Journal of evolutionary biology.

[10]  Mark E. Borrello,et al.  The rise, fall and resurrection of group selection. , 2005, Endeavour.

[11]  P D Taylor,et al.  OVERLAPPING GENERATIONS CAN PROMOTE ALTRUISTIC BEHAVIOR , 2000, Evolution; international journal of organic evolution.

[12]  J. Henrich Cultural group selection, coevolutionary processes and large-scale cooperation , 2004 .

[13]  C. Hauert,et al.  Synergy and discounting of cooperation in social dilemmas. , 2006, Journal of theoretical biology.

[14]  Ernst Fehr,et al.  Is Strong Reciprocity a Maladaptation? On the Evolutionary Foundations of Human Altruism , 2003 .

[15]  M. Nowak Five Rules for the Evolution of Cooperation , 2006, Science.

[16]  F. Marlowe,et al.  Anthropometric data indicate nutritional homogeneity in Hadza foragers of Tanzania , 2007, American journal of human biology : the official journal of the Human Biology Council.

[17]  Jordan Grafman,et al.  Functional Networks in Emotional Moral and Nonmoral Social Judgments , 2002, NeuroImage.

[18]  Fernando Santos-Granero Of fear and friendship: Amazonian sociality beyond kinship and affinity , 2007 .

[19]  M. Feldman,et al.  Gene-culture coevolution: models for the evolution of altruism with cultural transmission. , 1985, Proceedings of the National Academy of Sciences of the United States of America.

[20]  H. Gintis Strong reciprocity and human sociality. , 2000, Journal of theoretical biology.

[21]  R. C. Kelly The evolution of lethal intergroup violence , 2005, Proceedings of the National Academy of Sciences of the United States of America.

[22]  E. Smith,et al.  Risk and reciprocity in Meriam food sharing , 2002 .

[23]  Á. Pascual-Leone,et al.  Diminishing Reciprocal Fairness by Disrupting the Right Prefrontal Cortex , 2006, Science.

[24]  F. Waal,et al.  Good Natured: The Origins of Right and Wrong in Humans and Other Animals , 1996 .

[25]  P. Underhill,et al.  The dual origin and Siberian affinities of Native American Y chromosomes. , 2002, American journal of human genetics.

[26]  Derek E. Wildman,et al.  Implications of natural selection in shaping 99.4% nonsynonymous DNA identity between humans and chimpanzees: Enlarging genus Homo , 2003, Proceedings of the National Academy of Sciences of the United States of America.

[27]  Jana Schaich Borg,et al.  Consequences, Action, and Intention as Factors in Moral Judgments: An fMRI Investigation , 2006, Journal of Cognitive Neuroscience.

[28]  J. Henrich,et al.  Costly Punishment Across Human Societies , 2006, Science.

[29]  E. Sober,et al.  Summary of: ‘Unto Others. The evolution and psychology of unselfish behavior' , 1998 .

[30]  U. Fischbacher,et al.  The nature of human altruism , 2003, Nature.

[31]  J. Mitani,et al.  Demographic influences on the hunting behavior of chimpanzees. , 1999, American journal of physical anthropology.

[32]  J. Henrich,et al.  Friendship, cliquishness, and the emergence of cooperation. , 2006, Journal of theoretical biology.

[33]  L. Wilkinson,et al.  X‐linked imprinting: effects on brain and behaviour , 2006, BioEssays : news and reviews in molecular, cellular and developmental biology.

[34]  Herbert Gintis,et al.  The hitchhiker's guide to altruism: gene-culture coevolution, and the internalization of norms. , 2003, Journal of theoretical biology.

[35]  R. Boyd,et al.  The Puzzle of Human Sociality , 2006, Science.

[36]  Joseph Henrich,et al.  Culture, evolution and the puzzle of human cooperation , 2006, Cognitive Systems Research.

[37]  J. Benenson,et al.  Children's altruistic behavior in the dictator game , 2007 .

[38]  Robert L. Bettinger,et al.  Was Agriculture Impossible during the Pleistocene but Mandatory during the Holocene? A Climate Change Hypothesis , 2001, American Antiquity.

[39]  J. O'Doherty,et al.  Empathic neural responses are modulated by the perceived fairness of others , 2006, Nature.

[40]  M. Toro,et al.  Mutual benefit can promote the evolution of preferential interactions and in this way can lead to the evolution of true altruism. , 2004, Theoretical population biology.

[41]  Y. Toquenaga,et al.  Effects of information and group structure on evolution of altruism: analysis of two-score model by covariance and contextual analyses. , 2005, Journal of theoretical biology.

[42]  P D Taylor,et al.  Direct fitness or inclusive fitness: how shall we model kin selection? , 2007, Journal of evolutionary biology.

[43]  M. Nowak,et al.  Evolution of cooperation by multilevel selection. , 2006, Proceedings of the National Academy of Sciences of the United States of America.

[44]  Robert Plomin,et al.  Prosocial behavior from early to middle childhood: genetic and environmental influences on stability and change. , 2006, Developmental psychology.

[45]  E. Fehr,et al.  Sociology (communication arising (reply)): The puzzle of human cooperation , 2003, Nature.

[46]  J. Gelernter,et al.  Personality Traits of Agreeableness and Extraversion are Associated with ADH4 Variation , 2007, Biological Psychiatry.

[47]  Ivanei E. Bramati,et al.  The Neural Correlates of Moral Sensitivity: A Functional Magnetic Resonance Imaging Investigation of Basic and Moral Emotions , 2002, The Journal of Neuroscience.

[48]  S. Horvath,et al.  Conservation and evolution of gene coexpression networks in human and chimpanzee brains , 2006, Proceedings of the National Academy of Sciences.

[49]  S. Rose The rise of neurogenetic determinism , 1995, Nature.

[50]  Eizo Akiyama,et al.  Reputation and the evolution of cooperation in sizable groups , 2005, Proceedings of the Royal Society B: Biological Sciences.

[51]  D. Hay,et al.  Prosocial action in very early childhood. , 1999, Journal of child psychology and psychiatry, and allied disciplines.

[52]  M. Nowak,et al.  Evolution of indirect reciprocity , 2005, Nature.

[53]  G. Pagnoni,et al.  A Neural Basis for Social Cooperation , 2002, Neuron.

[54]  S. Rose Neurogenetic determinism and the new euphenics , 1998, BMJ.

[55]  B. Charlton A new science of health: salutology and the evolutionary perspective. , 1996, QJM : monthly journal of the Association of Physicians.

[56]  K. Slocombe,et al.  Fruit sharing between wild adult chimpanzees (Pan troglodytes schweinfurthii): a socially significant event? , 2005, American Journal of Primatology.

[57]  Dharol Tankersley,et al.  Altruism is associated with an increased neural response to agency , 2007, Nature Neuroscience.

[58]  R. Plomin Genes and behaviour. , 1995, Annals of medicine.

[59]  P. Richerson,et al.  The evolution of reciprocity in sizable groups. , 1988, Journal of theoretical biology.

[60]  Elizabeth Pennisi,et al.  How Did Cooperative Behavior Evolve? , 2005, Science.

[61]  R. Adolphs,et al.  Damage to the prefrontal cortex increases utilitarian moral judgements , 2007, Nature.

[62]  K. Zuberbühler,et al.  Leopard predation and primate evolution. , 2002, Journal of human evolution.

[63]  G. Robinson,et al.  Sociogenomics: social life in molecular terms , 2005, Nature Reviews Genetics.

[64]  J. Grafman,et al.  Human fronto–mesolimbic networks guide decisions about charitable donation , 2006, Proceedings of the National Academy of Sciences.

[65]  Sebastian Bonhoeffer,et al.  Evolution of cooperation by generalized reciprocity , 2005, Proceedings of the Royal Society B: Biological Sciences.

[66]  Diana C. Robertson,et al.  The neural processing of moral sensitivity to issues of justice and care , 2007, Neuropsychologia.

[67]  Rüdiger J Seitz,et al.  Functional modularity of the medial prefrontal cortex: involvement in human empathy. , 2006, Neuropsychology.

[68]  C. Stanford Chimpanzee Hunting Behavior , 1995 .

[69]  M. Tomasello,et al.  Altruistic Helping in Human Infants and Young Chimpanzees , 2006, Science.

[70]  E. Wilson,et al.  Eusociality: origin and consequences. , 2005, Proceedings of the National Academy of Sciences of the United States of America.

[71]  James H Fowler,et al.  Altruistic Punishment and the Origin of Cooperation , 2005, Proceedings of the National Academy of Sciences of the United States of America.

[72]  R. Baschetti The dawn of science-based moral reasoning. , 2007, Medical hypotheses.

[73]  G. Fink,et al.  Being with virtual others: Neural correlates of social interaction , 2006, Neuropsychologia.

[74]  P. McGuffin,et al.  Behaviour and genes , 1999 .

[75]  Andrew M. Colman,et al.  The puzzle of cooperation , 2006, Nature.

[76]  David K. Levine,et al.  The evolution of cooperation through imitation , 2002, Games Econ. Behav..