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Gene±Culture Coevolution 1 Gene±Culture Coevolution 1 Gene±Culture Coevolution Intermediate article Kevin N Laland, University of Cambridge, Cambridge, UK CONTENTS Introduction Do genes and culture coevolve? Evidence for transmitted culture Conclusion Types of cultural selection Evolution in species with a dynamic, socially trans- skills were transmitted from one generation to the mitted culture may be different from evolution in next, these simple artifacts represent the earliest other species. Population geneticists have pro- evidence for culture. In fact, comparative evidence posed the gene±culture coevolutionary approach for social learning in a variety of vertebrate species to describe the way in which cultural change may suggests that cultural transmission almost certainly drive a population's biological evolution. preceded Homo habilis by a considerable length of time. However, social learning in other animals is INTRODUCTION rarely stable enough to support traditions in which information accumulates from one generation to Many researchers have noted analogies between the next. For at least 2 million years our ancestors the processes of biological evolution and cultural have reliably inherited two kinds of information, change. For instance, both genes and culture are one encoded by genes, the other by culture. informational entities that are differentially trans- There is only one evolutionary approach to the mitted from one generation to the next. These simi- study of human behavior that takes up the chal- larities have led to the idea that culture evolves, lenge of understanding genetic and cultural evolu- and prompted the development of mathematical tion simultaneously by focusing directly on their models of cultural evolution. interaction. Gene±culture coevolutionary theory The main scientific approach to the study of (or dual inheritance theory), together with evolu- how culture evolves is a branch of theoretical tionary psychology and human behavioral ecology, population genetics, known variously as `cultural is one of three principal evolutionary approaches evolution', `gene±culture coevolution', or `dual in- that emerged in the aftermath of the human heritance' theory. This intellectual tradition has sociobiology debate (Smith, 2000). nothing in common with the nineteenth-century Conceptually, gene±culture coevolution is like a `cultural evolution' schools, which, based on an hybrid between memetics and evolutionary psych- erroneous view of evolution as progressive, set ology, although its methods are quite different, out to model stages of societal development. relying as they do on rigorous mathematical Rather, the population genetics approach regards theory. Like memeticists, gene±culture coevolution culture as an evolving pool of ideas, beliefs, values, enthusiasts treat culture as evolving learned know- and knowledge that is learned and socially trans- ledge. Like evolutionary psychologists, these re- mitted between individuals. Researchers focus on a searchers believe that the cultural knowledge single trait, such as a preference for drinking milk, individuals adopt may sometimes ± although cer- or for sons over daughters, and employ a rigorous tainly not always ± depend on their genetic consti- mathematical approach to describe how the cul- tution. Moreover, selection acting on the genetic tural trait changes over time, sometimes coevolving system is commonly generated or modified by the with genetic variation. Where the cultural entity is spread of cultural information. For gene±culture a discrete package, it has much in common with coevolutionary theorists, the `leash' that ties culture Richard Dawkins' idea of the `meme', defined as a to genes tugs both ways. The advent of culture was cultural analogue of the gene (Dawkins, 1976). a precipitating evolutionary milestone, generating Stone tools appear in the archeological record selection that favored a reorganization of the approximately two and a half million years ago. human brain, and leaving it specialized to acquire, If, as is widely believed, lithic technologies and store, and use cultural information. It was culture, 2 Gene±Culture Coevolution loosely guided by genes, that allowed humans the to the natural selection of genes. Gene±culture adaptive flexibility to colonize the world. coevolution exhibits a concern for nonadaptive The quantitative study of gene±culture coevolu- and even maladaptive outcomes of the evolution- tion began in 1976, when two population geneti- ary process. This stance continues both to surprise cists, Luca Cavalli-Sforza and Marc Feldman of and confuse outside observers used to characteriz- Stanford University, published the first simple ing all these evolutionary approaches as `sociobiol- dynamic models with both genetic and cultural ogy'. However, the rigorous theoretical approach inheritance. The fundamental innovation that has led to little experimentation or other forms Cavalli-Sforza and Feldman instigated was that, in of empirical work, and this school remains the addition to modeling the differential transmission prerogative of a comparatively small band of of genes from one generation to the next, they in- workers. corporated cultural information into the analysis, The emerging body of theory has developed in a allowing the evolution of the two systems to be variety of ways. One class of models investigates mutually dependent. However, one curious feature the inheritance of behavioral and personality traits, of the history of gene±culture coevolution is that extending traditional models by incorporating a both archetypal sociobiologists and some of their transmitted cultural component into the analysis. most severe critics almost simultaneously recog- Other models address general questions about the nized the importance of gene±culture interactions, adaptive advantages of learning and culture. More with each starting to develop methods to address recently, these methods have been applied to ad- the problem. By the late 1970s, Charles Lumsden dress specific cases in which there is an interaction and Edward Wilson at Harvard University were between cultural knowledge and genetic variation engaged in a race with Cavalli-Sforza and Feldman that influences its prevalence. These include the to produced the first book on this topic. While evolution of language and of handedness, an an- Lumsden's and Wilson's Genes, Mind and Culture alysis of changes in the genetic sex ratio in the face was published first (Lumsden and Wilson, 1981), it of sex-biased parental investment, the spread of was not well regarded (Maynard-Smith and agriculture, the coevolution of hereditary deafness Warren, 1981). In contrast, Cavalli-Sforza's and and sign language, the emergence of incest taboos, Feldman's more cautious tome Cultural Transmis- and an exploration of how cultural niche construc- sion and Evolution (Cavalli-Sforza and Feldman, tion affected human evolution (see Feldman and 1981) was much better received. Laland, 1996). Together with many co-workers, Cavalli-Sforza As the rules of cultural transmission are usually and Feldman gradually built up an impressive different from those of genetic transmission, simi- body of mathematical theory exploring the pro- lar selective regimes may result in very different cesses of cultural change and interaction between equilibria. A good example of this is provided by genes and culture. Frequently they took advantage the hypothesis of Boyd and Richerson (1985) that of the parallels between the spread of a gene and group selection can act on cultural variation. The the diffusion of a cultural innovation to borrow or theoretical argument against group selection is adapt established models from population genet- based on models which assume genetic inherit- ics. Drawn by the ongoing sociobiology debate, ance, and the criticisms may not hold for culturally other mathematically minded researchers joined transmitted traits. When individuals adopt the be- the fray, most notably anthropologists Rob Boyd havior of the majority a conformist transmission is and Peter Richerson, whose book Culture and the generated (Boyd and Richerson, 1985). As a result Evolutionary Process introduced a variety of novel of its frequency dependence, conformist transmis- theoretical methods and stimulating ideas (Boyd sion can act to amplify differences in the frequency and Richerson, 1985). Gradually a consensus of cultural traits in different subpopulations, but as to the most appropriate methods for tackling reduce variance within groups. Boyd and Richer- gene±culture interactions began to emerge, which son showed that one of the by-products of a con- today forms the basis of modern coevolutionary formist bias is an increase in the strength of the theory. group selection of cultural variation so that it may The technical and explicitly mathematical nature be a strong force relative to forces acting within of modern gene±culture coevolution is one of sev- groups, such as natural selection. Since selection eral features that distinguishes this perspective between groups may favor beliefs and attitudes from alternatives such as evolutionary psychology. that benefit the group at the expense of the individ- A second is the incorporation into analyses of a ual, Boyd and Richerson's theory provides a new variety of genetic and cultural processes in addition explanation for human cooperation. Gene±Culture Coevolution 3 EVIDENCE FOR TRANSMITTED different from other aspects of the environment is CULTURE the knowledge passed between individuals. Cul-
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