David Wilson

In both biology and the human sciences, social groups are sometimes treated as adaptive units whose organization cannot be reduced to individual interactions. This group-level view is opposed by a more individualistic one that treats social organization as a byproduct of self-interest. According to biologists, group-level adaptations can evolve only by a process of natural selection at the group level. Most biologists rejected group selection as an important evolutionary force during the 1960s and 1970s but a positive literature began to grow during the 1970s and is rapidly expanding today. We review this recent literature and its implications for human evolutionary biology. We show that the rejection of group selection was based on a misplaced emphasis on genes as "replicators" which is in fact irrelevant to the question of whether groups can be like individuals in their functional organization. The fundamental question is whether social groups and other higher-level entities can be "vehicles" of selection. When this elementary fact is recognized, group selection emerges as an important force in nature and what seem to be competing theories, such as kin selection and reciprocity, reappear as special cases of group selection. The result is a unified theory of natural selection that operates on a nested hierarchy of units. The vehicle-based theory makes it clear that group selection is an important force to consider in human evolution. Humans can facultatively span the full range from self-interested individuals to "organs" of group-level "organisms." Human behavior not only reflects the balance between levels of selection but it can also alter the balance through the construction of social structures that have the effect of reducing fitness differences within groups, concentrating natural selection (and functional organization) at the group level. These social structures and the cognitive abilities that produce them allow group selection to be important even among large groups of unrelated individuals.

Group selection is increasingly being viewed as an important force in human evolution. This paper examines the views of R.D. Alexander, one of the most influential thinkers about human behavior from an evolutionary perspective, on the subject of group selection. Alexander's general conception of evolution is based on the gene-centered approach of G.C. Williams, but he has also emphasized a potential role for group selection in the evolution of individual genomes and in human evolution. Alexander's views are internally inconsistent and underestimate the importance of group selection. Specific themes that Alexander has developed in his account of human evolution are important but are best understood within the framework of multilevel selection theory. From this perspective, Alexander's views on moral systems are not the radical departure from conventional views that he claims, but remain radical in another way more compatible with conventional views.

We have two main objections to Kerr and Godfrey-Smith's (2002) meticulous analysis. First, they misunderstand the position we took in Unto Others – we do not claim that individual-level statements about the evolution of altruism are always unexplanatory and always fail to capture causal relationships. Second, Kerr and Godfrey-Smith characterize the individual and the multi-level perspectives in terms of different sets of parameters. In particular, they do not allow the multi-level perspective to use the individual fitness parameters i and i. We don't see why the multi-level perspective prevents one from thinking in these terms. Kerr and Godfrey-Smith's argument that Uyenoyama and Feldman's (1980, 1992) definition of altruism belongs more to the individualist perspective than it does to the multi-level perspective is an artifact of their choice of parameters; the same point applies to their argument about the individualism inherent in the idea of Class I and Class II fitness structures.

We address the following issues raised by the commentators of our target article and book: the problem of multiple perspectives; how to define group selection; distinguishing between the concepts of altruism and organism; genetic versus cultural group selection; the dark side of group selection; the relationship between psychological and evolutionary altruism; the question of whether the psychological questions can be answered; psychological experiments. We thank the contributors for their commentaries, which provide a diverse agenda for future study of evolution and morality. Our response will follow the organization of our book, distinguishing between evolutionary issues that concern fitness effects and psychological issues that concern motives.

The hypothesis of group selection fell victim to a seemingly devastating critique in 1960s evolutionary biology. In Unto Others, we argue to the contrary, that group selection is a conceptually coherent and empirically well documented cause of evolution. We suggest, in addition, that it has been especially important in human evolution. In the second part of Unto Others, we consider the issue of psychological egoism and altruism -- do human beings have ultimate motives concerning the well-being of others? We argue that previous psychological and philosophical work on this question has been inconclusive. We propose an evolutionary argument for the claim that human beings have altruistic ultimate motives.

The substance of the commentaries, however, reveals considerable disagreement about how UO conceptualizes the idea of group selection. Dennett describes the issues as “mind-twistingly elusive and slippery” and hints that it is mere hype to say that group selection has been revived. Barrett and Godfrey-Smith discuss the problem of multiple perspectives at length and claim that we are too liberal in our definition of groups. We believe that these criticisms obscure the simplicity of the basic question that group selection theory is designed to answer and the historical continuity of that theory, from Darwin to the present.

Altruism has both an evolutionary and a psychological meaning. As the term is used in evolutionary theory, a trait is deemed altruistic if it reduces the fitness of the actor and enhances the fitness of someone else. In its psychological sense, the thesis that we have altruistic ultimate motives asserts that we care about the welfare of others, not just as a means of enhancing our own well-being, but as an end in itself. In Unto Others (hereafter UO), we consider both evolutionary altruism (Part I) and psychological altruism (Part II) from an evolutionary perspective. We propose a multilevel selection theory to explain how altruism can evolve from group selection, based on Darwin’s solution to the problem. Our presentation of multilevel selection theory expands the range of topics beyond altruism in other respects as well. In Part I1 of UO, we evaluate the arguments, both conceptual and empirical, that philosophers and psychologists have advanced in an attempt to choose between psychological egoism and motivational pluralism. We conclude that these arguments are inconclusive.

In both biology and the human sciences, social groups are sometimes treated as adaptive units whose organization cannot be reduced to individual interactions. This group-level view is opposed by a more individualistic one that treats social organization as a byproduct of self-interest. According to biologists, group-level adaptations can evolve only by a process of natural selection at the group level. Most biologists rejected group selection as an important evolutionary force during the 1960s and 1970s but a positive literature began to grow during the 1970s and is rapidly expanding today. We review this recent literature and its implications for human evolutionary biology. We show that the rejection of group selection was based on a misplaced emphasis on genes as “replicators” which is in fact irrelevant to the question of whether groups can be like individuals in their functional organization. The fundamental question is whether social groups and other higher-level entities can be “vehicles” of selection. When this elementary fact is recognized, group selection emerges as an important force in nature and what seem to be competing theories, such as kin selection and reciprocity, reappear as special cases of group selection. The result is a unified theory of natural selection that operates on a nested hierarchy of units.The vehicle-based theory makes it clear that group selection is an important force to consider in human evolution. Humans can facultatively span the full range from self-interested individuals to “organs” of group-level “organisms.” Human behavior not only reflects the balance between levels of selection but it can also alter the balance through the construction of social structures that have the effect of reducing fitness differences within groups, concentrating natural selection (and functional organization) at the group level. These social structures and the cognitive abilities that produce them allow group selection to be important even among large groups of unrelated individuals.