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Gene-Culture Coevolution of Complex Social Behavior: Human Altruism

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Gene-Culture Coevolution of Complex Social Behavior: Human Altruism Proc. Natl. Acad. Sci. USA Vol. 83, pp. 7340-7343, October 1986 Evolution Gene-culture coevolution of complex social behavior: Human altruism and mate choice (behavior genetics/developmental psychology/epigenetic rules/gene-culture trasnuission/sociobiology) J. PHILIPPE RUSHTON*, CHRISTINE H. LITTLEFIELDt, AND CHARLES J. LUMSDENt *Department of Psychology, University of Western Ontario, London, ON, N6A 5C2 Canada; and tDepartment of Medicine, University of Toronto, Toronto, ON, M5S 1A8 Canada Communicated by E. 0. Wilson, June 19, 1986 ABSTRACT The hypothesis is examined that genes bias learner (e.g., refs. 8, 9, and 11). For example, most models the development of complex social behavior in one direction of cultural transmission within the family (i.e., vertical, from over alternatives. Studies of altruism and political attitudes in parent to child, and horizontal, from sibling to sibling) imply twins estimate that =50% of the variance is associated with that siblings will resemble each other, over and above shared direct genetic inheritance, virtually 0% with the twin's com- genes, as a result of a common family environment. Gene- mon family environment, and the remainder with each twin's culture theory, in contrast, leads to the expectation that specific environment. Studies of human marriages show that siblings will differ from each other in part because their spouses choose each other on the basis of similarity, assorting nonshared genes incline them to acquire patterns of behavior on the most genetically influenced of a set of homogeneous best fitting their particular genotype (gene-culture transmis- attributes. These data imply a genetic canalization of social sion). While it may seem intuitively correct to assume that influences such that, within the constraints allowed by the total common family environment shapes individual development, spectrum of cultural alternatives, people create environments consideration of data reveals quite a different set of relation- maximally compatible with their genotypes. ships. Behavior genetic model fitting techniques provide increas- Increasing efforts are being made to understand the relation- ingly powerful tests of alternative hypotheses about the ship between human cultural and genetic transmission. As genetic and social influences on family resemblances (12-15). Lumsden and Wilson (1) argued when presenting the theory One useful design involves the comparison of monozygotic of gene-culture coevolution, neither purely genetic nor and dizygotic twins reared together. While critics have purely cultural modes of transmission are likely to be evo- argued that the twin method is invalid, detailed empirical lutionarily stable. Instead, they provided a model to integrate work demonstrates the critiques to be of limited importance genetic and environmental influences into a reciprocating (16, 17). In a twin study, the raw data are the between- and circuit such that epigenetic rules guide individual develop- within-pairs variances and covariances. The between-pairs ment in one direction over alternatives (termed gene-culture mean squares reflect both pair resemblances and pair differ- transmission). Innate biases to learn one pattern of cultural ences, and the within-pairs mean squares, pair differences. information over another provide a particularly likely mech- The genetic models are fitted to these mean squares. The total anism by which dual inheritance can occur. phenotypic variance can be partitioned into the following Most if not all ofthe components of cognitive development three sources: V(G), additive genetic effects; V(CE), com- examined to date have supported the gene-culture model of mon environmental influences that affect both twins equally; transmission (1-7). Corresponding data on personality and and V(SE), specific environmental influences that affect each social development have been attended to far less. Here we twin individually. This latter is a residual term that is review research in support of the hypothesis that epigenetic comprised of many sources, including measurement error rules bias the development of complex social behavior, and certain kinds of interaction between genotypes and choosing as illustrative examples, altruism and mate choice. environments. Thus, the total phenotypic variance is parti- In so doing, we contrast Lumsden and coworkers' (1-7) tioned as V(G) + V(CE) + V(SE). approach with models of cultural transmission hypothesized Using such a design with 573 adult monozygotic and by Cavalli-Sforza, Feldman, and their colleagues (8-10). dizygotic twin pairs, Rushton et al. (18) examined the cultural While the latter authors incorporated the phrase "gene-cul- and genetic inheritance of individual differences in altruism ture coevolution" in a study of the transmission of altruism and aggression. Components of these traits were measured (10), they have generally failed to adopt the concomitant by paper and pencil questionnaires in which the 1146 respon- prescription that epigenetic rules bias individual develop- dents endorsed items measuring their self-reported altruistic ment. As a result, much of their perspective is discordant behavior, empathy, nurturance, aggressiveness, and as- with knowledge about social learning in families. By contrast, sertiveness. Maximum-likelihood model-fitting estimation gene-culture coevolutionary formulations, in the sense orig- procedures revealed '50% of the variance on each scale to inally intended by Lumsden and Wilson (1), appear to be be associated with genetic effects, virtually 0% with the highly compatible with these same facts. twin's common environment, and the remaining 50% with each twin's specific environment and/or error associated Epigenetic Rules in Social Development with the test. Correcting for the unreliability in the tests raised the heritabilities to -60% and reduced the specific Extensive theorizing in both the evolutionary and social environment variance to 40%. A summary of the results is sciences errs in not taking into account that social learning is presented in Table 1. dependent upon the innate capacities and biases of the These data not only signify a strong association of genetic factors with the characteristics in question but also indicate The publication costs of this article were defrayed in part by page charge a negligible influence of the twin's shared environment. payment. This article must therefore be hereby marked "advertisement" Rather, the distinct experiences of the individual account for in accordance with 18 U.S.C. §1734 solely to indicate this fact. almost all the environmental variance. Approaches such as 7340 Evolution:EsProc.Rushton et al. Natl. Acad. Sci. USA 83 (1986) 7341 Table 1. Estimates of variance components and estimates vertical cultural transmission ofthe type proposed by Cavalli- corrected for unreliability from a biometrical analysis of Sforza et al. (9) fails to influence social attitudes. They altruism, empathy, nurturance, aggressiveness, and received questionnaires measuring attitudes toward such assertiveness questionnaires from 573 adult twin pairs issues as the death penalty from 4600 pairs of adult twins. Common Specific Other sources provided estimates ofassortative mating. Path Additive environ- environ- analysis indicated a very poor fit when the genetic contribu- genetic mental mental tion to the observed variation was set to zero. On the other variance variance variance hand, models deleting vertical cultural inheritance gave an extremely good fit when allowances were made for assorta- Trait %E %EC %E %EC %E %EC tive mating. The cogency of the estimation techniques, Altruism 51 60 2 2 47 38 sample sizes, and questionnaire validities employed were Empathy 51 65 0 0 49 35 such that the authors were able to predict the correlations Nurturance 43 60 1 1 56 39 that would be expected in other relationships: e.g., 0.00 Aggressiveness 39 54 0 0 61 46 between foster parent and adult foster child; 0.52 between Assertiveness 53 69 0 0 47 31 parents and children; and 0.62 for separated monozygotic %E, variance component; %EC, estimates corrected for unreli- twins. The results led the authors to conclude that: ... Data has been from Rushton et 18. geneticists and social scientists [might] have misconceived ability. adapted al., ref. the role of cultual inheritance and that individuals acquire little from their social environment that is incompatible with those by Cavalli-Sforza and Feldman (8, 9) imply a substan- their genotype" (ref. 24, p. 4368). tial effect for common environment. Clearly, this assumption One feature of developmental research highlighted by a is incompatible with the empirical data reported in Table 1. gene-culture coevolutionary perspective, but underex- The account provided by a mechanism of gene-culture amined to date, is the role epigenetic rules play in modulating transmission, on the other hand, predicting that social de- human life-history phenomena. Familial concordances show velopment will be guided by epigenetic rules that incline that genetic mechanisms sequence such trajectories as individuals to particular learning experiences is directly growth rates in height and mental development; age of onset compatible with the large specific environment term shown. of puberty, menopause, and first sexual experience; family The discovery that common family environment plays a size; fecundity; a variety of health-related phenomena in- very limited role in social development (even for traits that cluding degenerative diseases
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