Although the field of bioethics usually emphasizes ethical dilemmas arising from contemporary biomedical research, at another level the foundation of ethical judgments can be explored in the light of evolutionary biology. Two scientific approaches illuminate the relationships between human nature, social environments, and standards of ethical judgment: first, ethology and the observational study of nonhuman primates; second, evolutionary theory and new developments in the understanding of natural selection. Ethology shows that humans, like the species most closely (...) related to us, are naturally both cooperative and competitive. Evolutionary theory can "explain" cooperation at diverse levels of social organization , suggesting which forms of social cooperation are to be expected in specific cases. By focusing on the emotions underlying the sense of justice, moreover, an evolutionary approach can provide a more realistic assessment of the motivations and intentions underlying ethical controversies. Because individuals inevitably have competing claims as well as competing moral obligations, evolutionary bioethics thus teaches the need for balance and judgment. Emotions elicited by the all-too-human tendency to engage in moralistic aggression need to be balanced by reasoned judgment of the consequences of our own actions. The ethicist, no more or less than any other human, needs to be aware of the risks of deceptions and self-deception. Or, to put it more simply, an evolutionary bioethics leads back to the humility of the Socratic injunction: know thyself. Im allgemein üblichen Sinne befaßt sich Bioethik meist mit ethischen Problemstellungen, die durch die Fortschritte der biologischen Forschung hervorgerufen werden. Unter einem anderen Blickwinkel kann sie aber auch die Grundlage ethischer Urteile im Licht der Entwicklungsbiologie untersuchen. Zwei Wissenschaftszweige befassen sich mit dem Verhältnis zwischen der menschlichen Natur, dem sozialen Umfeld und den Standards für ethische Urteile; erstens, die Ethologie und Verhaltensstudien an nichtmenschlichen Primaten; zweitens, die Evolutionstheorie und neue Entwicklungen beim Verständnis der natürlichen Selektion. Die Ethologie zeigt, daß Menschen genauso wie die Tiere, die ihnen am nächsten verwandt sind, von Natur aus sowohl zur Kooperation als auch zur Konkurrenz miteinander neigen. Die Evolutionstheorie kann die verschiedenen Weisen von Kooperation auf den verschiedenen Ebenen sozialer Organisation "erklären" und dabei Hypothesen aufstellen, welche Formen sozialer Zusammenarbeit in spezifischen Fällen erwartet werden können. Soweit sie sich mit den Emotionen befaßt, die dem Sinn für Gerechtigkeit zugrunde liegen, kann die Evolutionstheorie darüber hinaus eine realistischere Einschätzung der Beweggründe und Intentionen vermitteln, die hinter ethischen Kontroversen stehen. Weil Individuen unvermeidlich konkurrierende Ansprüche, aber auch konkurrierende moralische Verpflichtungen haben, zeigt die evolutionäre Bioethik das Bedürfnis nach Ausgleich und Schlichtung. Die Emotionen, die durch die allzu menschliche Neigung zu Aggression im Namen der Moral hervorgerufen werden, rufen nach Ausgleich durch begründete Urteile über die Folgen unserer eigenen Handlungen. Der Ethiker muß sich, nicht mehr und nicht weniger als jeder andere Mensch, des Risikos von Täuschung und Selbsttäuschung bewußt sein. Oder, um es einfacher auszudrücken, evolutionäre Ethik führt zurück zu der Bescheidenheit der Sokratischen Forderung: Erkenne dich selbst. (shrink)

The Nature of Selection is a straightforward, self-contained introduction to philosophical and biological problems in evolutionary theory. It presents a powerful analysis of the evolutionary concepts of natural selection, fitness, and adaptation and clarifies controversial issues concerning altruism, group selection, and the idea that organisms are survival machines built for the good of the genes that inhabit them. "Sober's is the answering philosophical voice, the voice of a first-rate philosopher and a knowledgeable student (...) of contemporary evolutionary theory. His book merits broad attention among both communities. It should also inspire others to continue the conversation."-Philip Kitcher, Nature "Elliott Sober has made extraordinarily important contributions to our understanding of biological problems in evolutionary biology and causality. The Nature of Selection is a major contribution to understanding epistemological problems in evolutionary theory. I predict that it will have a long lasting place in the literature."-Richard C. Lewontin. (shrink)

This paper discusses the relations between three forms of altruism: behavioural, evolutionary and motivational. Altruism in a behavioural sense is an act that benefits another person. It can range from volunteering to a charity and helping a neighbour, to giving money to a non-profit organisation or donating blood. People often dedicate their material and nonmaterial resources for the benefit of others to gain psychological, social and material benefits for themselves. Thus, their altruistic acts are driven by egoistic (...) motivation. Also, the final goal of an altruistic act may be the increase in the welfare of a group or adherence to a certain moral principle or a social norm. However, at least sometimes, the welfare of others is the ultimate goal of our actions, when our altruistic acts are performed from altruistic motivation. In evolutionary sense, altruism means the sacrifice of reproductive success for the benefit of other organisms. According to evolutionary theories, behaviour which promotes the reproductive success of the receiver at the cost of the actor is favoured by natural selection, because it is either beneficial for the altruist in the long run, or for his genes, or for the group he belongs to. However, altruism among people emerges as a distinctly human combination of innate and learned behaviours. Not only do we benefit the members of our own group, but we are capable of transcending our tribalistic instincts and putting the benefit of strangers at our own personal expense as our ultimate goal. (shrink)

Group-structured and neighbor-structured populations are compared, especially in relation to multilevel selection theory and evolutionary transitions. I argue that purely neighborstructured populations, which can feature the evolution of altruism, are not properly described in multilevel terms. The ability to “gestalt switch” between individualist and multilevel frameworks is then linked to the investigation of “major transitions” in evolution. Some explanatory concepts are naturally linked to one framework or the other, but a full understanding is best achieved via (...) the use of both. (shrink)

Drawing on an idea proposed by Darwin, it has recently been hypothesised that violent intergroup conflict might have played a substantial role in the evolution of human cooperativeness and altruism. The central notion of this argument, dubbed ‘parochial altruism’, is that the two genetic or cultural traits, aggressiveness against out-groups and cooperativeness towards the in-group, including self-sacrificial altruistic behaviour, might have coevolved in humans. This review assesses the explanatory power of current theories of ‘parochial altruism’. After a (...) brief synopsis of the existing literature, two pitfalls in the interpretation of the most widely used models are discussed: potential direct benefits and high relatedness between group members implicitly induced by assumptions about conflict structure and frequency. Then, a number of simplifying assumptions made in the construction of these models are pointed out which currently limit their explanatory power. Next, relevant empirical evidence from several disciplines which could guide future theoretical extensions is reviewed. Finally, selected alternative accounts of evolutionary links between intergroup conflict and intragroup cooperation are briefly discussed which could be integrated with parochial altruism in the future. (shrink)

This essay addresses recent claims about the compatibility of the sociobiological theory of reciprocal altruism with standard Western formulations of the Golden Rule. Derek Parfit claims that the theory of reciprocal altruism teaches us to be “reciprocal altruists,” who benefit only those people from whom we can reasonably expect benefits in the future. The Golden Rule, on the other hand, teaches us to benefit anyone regardless of their intention or ability to return the favor, or as (...) Parfit puts it, the Golden Rule teaches us to be “suckers.” I argue that this distinction is founded on a misconception of the nature of the theory of reciprocal altruism, which is sociobiological as opposed to moral, and that this distinction accordingly confuses is with ought. Sociobiological theories may explain underlying psychological motivations in individuals (and perhaps even in populations), but these theories do not prescribe any sort of moral behavior. Furthermore, the theory of reciprocal altruism does not imply mental states of which agents are aware. The unconscious motivations assumed by this theory are in fact compatible with certain formulations of the Golden Rule; I will accordingly argue for the view that certain words with moral content related to the Golden Rule—such as “altruism” and “selfishness”—exist only insofar as they are social tools, which can further the self-interests of an individual in any group. (shrink)

Our aim in this chapter is to draw lessons from current theory on the evolution of human cooperation for the management of contemporary commons. Evolutionary theorists have long been interested in cooperation but social scientists have documented patterns of cooperation in humans that present unusual problems for conventional evolutionary theory (and for rational choice explanations as well). Humans often cooperate with nonrelatives and are prone to cooperate in one-shot games. Cooperation is quite dependent on social institutions. (...) We believe that this last fact is the critical clue to understanding human cooperation. Models of cultural evolution suggest that group selection is a more potent force on culture than on genes. Evolutionary theory is in essence a theory of preferences in terms of rational actor theory and is thus complementary to the bounded rational choice models that underpin so much theorizing in the social sciences. Thus, the theory suggests a source for prosocial impulses, and leads to predictions about the limits of human altruism and constraints likely to be imposed upon the evolution of social institutions. We also consider the dynamics of genes as they coevolved with increasingly sophisticated cultural institutions over the long course of human evolution in the Pleistocene. We hypothesize that the long exposure of human populations to group selected cultural norms and preferences is likely to have resulted in an innate psychology adapted to living in egalitarian, cooperative societies of a few hundred to a few thousand people. We call this the tribal social instincts hypothesis. The evolution of complex societies in the past few thousand years constitutes a series of natural experiments that test this hypothesis. If it is correct, the institutions of complex societies must somehow take advantage of the prosocial elements of the tribal instincts while finessing the problem that the tribal social instincts are ill adapted to life in large, hierarchical, inegalitarian societies with extensive dominance of subordinates by elites.. (shrink)

Our aim in this chapter is to draw lessons from current theory on the evolution of human cooperation for the management of contemporary commons. Evolutionary theorists have long been interested in cooperation but social scientists have documented patterns of cooperation in humans that present unusual problems for conventional evolutionary theory (and for rational choice explanations as well). Humans often cooperate with nonrelatives and are prone to cooperate in one-shot games. Cooperation is quite dependent on social institutions. (...) We believe that this last fact is the critical clue to understanding human cooperation. Models of cultural evolution suggest that group selection is a more potent force on culture than on genes. Evolutionary theory is in essence a theory of preferences in terms of rational actor theory and is thus complementary to the bounded rational choice models that underpin so much theorizing in the social sciences. Thus, the theory suggests a source for prosocial impulses, and leads to predictions about the limits of human altruism and constraints likely to be imposed upon the evolution of social institutions. We also consider the dynamics of genes as they coevolved with increasingly sophisticated cultural institutions over the long course of human evolution in the Pleistocene. We hypothesize that the long exposure of human populations to group selected cultural norms and preferences is likely to have resulted in an innate psychology adapted to living in egalitarian, cooperative societies of a few hundred to a few thousand people. We call this the tribal social instincts hypothesis. The evolution of complex societies in the past few thousand years constitutes a series of natural experiments that test this hypothesis. If it is correct, the institutions of complex societies must somehow take advantage of the prosocial elements of the tribal instincts while finessing the problem that the tribal social instincts are ill adapted to life in large, hierarchical, inegalitarian societies with extensive dominance of subordinates by elites.. (shrink)

The main thesis defended in this dissertation is that culture makes a difference. It is argued that this is so because culture evolves just as species do. It is maintained that if one views culture as an evolving system, then this has some fundamental conceptual, ontological, and explanatory implications. Specifically, construing culture as an evolving system leads one to explain cultural phenomena and behaviors differently from alternative theories. Hence, this thesis is devoted to laying the foundations for a satisfactory (...) class='Hi'>theory of cultural evolution. Chapter 1 explains the conditions under which culture can be seen as evolving and attempts to clarify the structure of a theory of cultural evolution. Chapter 2 stresses the kind of explanatory work that a theory of cultural evolution is capable of doing by developing an approach to explanation called explanatory pluralism. Chapter 3 argues that a theory of cultural evolution must include an account of cultural fitness. A multidimensional approach to cultural fitness is put forth and the function and explanatory utility of this approach are stressed. Then chapter 4 addresses the question of whether there is such a thing as cultural selection. It is argued that though culture is no doubt selective, there is not one selection mechanism driving cultural evolution. Rather, it is claimed that the evolution of culture has been the evolution of a multitude of selection mechanisms. Chapter 5 concludes by pointing out another interesting consequence of culture, namely, that it may favor the evolution of altruism and maladaptive traits. (shrink)

In their book Unto Others, Sober and Wilson argue that various evolutionary considerations (based on the logic of natural selection) lend support to the truth of psychological altruism. However, recently, Stephen Stich has raised a number of challenges to their reasoning: in particular, he claims that three out of the four evolutionary arguments they give are internally unconvincing, and that the one that is initially plausible fails to take into account recent findings from cognitive science and (...) thus leaves open a number of egoistic responses. These challenges make it necessary to reassess the plausibility of Sober & Wilson’s evolutionary account—which is what I aim to do in this paper. In particular, I try to show that, as a matter of fact, Sober & Wilson’s case remains compelling, as some of Stich’s concerns rest on a confusion, and those that do not are not sufficiently strong to establish all the conclusions he is after. The upshot is that no reason has been given to abandon the view that evolutionary theory has advanced the debate surrounding psychological altruism. (shrink)

Runaway social selection resulting from partner choice may have shaped aspects of human cooperation and complex sociality that are otherwise hard to account for. Social selection is the subtype of natural selection that results from the social behaviors of other individuals. Competition to be chosen as a social partner can, like competition to be chosen as a mate, result in runaway selection that shapes extreme traits. People prefer partners who display valuable resources and bestow them selectively (...) on close partners. The resulting phenotypic covariance between displays and preferences gives fitness advantages to both, creating runaway selection that could shape a whole suite of prosocial traits including altruism, moral capacities, empathy, and theory of mind. Even though they give a net fitness benefit, traits at the endpoint of runaway social selection can have substantial deleterious effects on other traits such as viability, ability to accumulate resources, or vulnerability to mental disorders. Social selection forces arising from self-interested partner choices may be an invisible hand that shaped capacities for commitment, altruism, and other prosocial capacities of the human social brain. (shrink)

Consider the paradox of altruism: the existence of truly altruistic behaviors is difficult to reconcile with an evolutionary theory which holds that natural selection operates only on individuals, since in that case individuals should be unwilling to sacrifice their own fitness for the sake of others. Evolutionists have frequently turned to the hypothesis of group selection to explain the existence of altruism; but, even setting aside difficulties about understanding the relationship between altruistic behaviors and (...) morality, group selection cannot explain the evolution of morality, since morality is a one-group phenomenon and group selection is a many-group phenomenon. After spelling out just what the problem is, this paper discusses several ways out and concludes by offering suggestions why one seems best. (shrink)

Altruism is a central concept in evolutionary biology. Evolutionary biologists still disagree about its meaning (E.O. Wilson 2005; Fletcher et al. 2006; D.S. Wilson 2008; Foster et al. 2006a, b; West et al. 2007a, 2008). Semantic disagreement appears to be quite robust and not easily overcome by attempts at clarification, suggesting that substantive conceptual issues lurk in the background. Briefly, group selection theorists define altruism as any trait that makes altruists losers to selfish traits within (...) groups, and makes groups of altruists fitter than groups of non-altruists. Inclusive fitness theorists reject a definition based on within- and between-group fitness. Traits are altruistic only if they cause a direct and absolute fitness loss to the donor. The latter definition is more restrictive and rejects as cases of altruism behaviors that are accepted by the former. Fletcher and Doebeli (2009) recently proposed a simple, direct and individually based fitness approach, which they claim returns to first principles: carriers of the genotype of interest “must, on average, end up with more net direct fitness benefits than average population members.” This seductively simple proposal uses the concept of assortment to explain how diverse kinds of altruists end up on average with more net fitness than their non-altruistic rivals. In this paper I shall argue that their approach implies a new concept of altruism that contrasts with and improves on the concept of the inclusive fitness approach. (shrink)

Recent years have seen a renewed debate over the importance of groupselection, especially as it relates to the evolution of altruism. Onefeature of this debate has been disagreement over which kinds ofprocesses should be described in terms of selection at multiple levels,within and between groups. Adapting some earlier discussions, we presenta mathematical framework that can be used to explore the exactrelationships between evolutionary models that do, and those that donot, explicitly recognize biological groups as fitness-bearing entities.We show (...) a fundamental set of mathematical equivalences between these twokinds of models, one of which applies a form of multi-level selectiontheory and the other being a form of ``individualism.'' However, we alsoargue that each type of model can have heuristic advantages over theother. Indeed, it can be positively useful to engage in a kind ofback-and-forth switching between two different perspectives on theevolutionary role of groups. So the position we defend is a``gestalt-switching pluralism.''. (shrink)

ABSTRACT:In this article, we describe evolutionary psychology and its potential contribution to business ethics research. After summarizing evolutionary theory and natural selection, we specifically address the use of evolutionary concepts in psychology in order to offer alternative explanations of behavior relevant to business ethics, such as social exchange, cooperation, altruism, and reciprocity. Our position is that individuals, groups, and organizations all are affected by similar natural, evolutionary processes, such that evolutionary psychology is (...) applicable at multiple levels of analysis (e.g., individual and group). We introduce a variety of experiments and instruments employed by evolutionary psychologists to illustrate how ethics-relevant cultural norms and practices evolve and are regulated, and to raise the prospect that these experiments and instruments can be useful in future business ethics research. (shrink)

The development of modern evolutionary ethics began shortly after the publication of Charles Darwin's On the Origin of Species by Natural Selection. Early discussions were plagued by several problems. First, evolutionary ethical explanations were dependent on group-selection accounts of social behavior (especially the explanation of altruism). Second, they seem to violate the philosophical principle that “ought” statements cannot be derived from “is” statements alone (values cannot be derivedfrom facts alone). Third, evolutionary ethics appeared to (...) be biologically deterministic, deemed incompatible with the free will required for ethics to be possible. Fourth, social policies based on evolutionary theory (for example, eugenics in the early part of this century) seemed patently unethical. Sociobiology (which coalesced as a field of study with Edward O. Wilson's Sociobiology: The New Synthesis, 1975) addressed several of these problems and provided a rich framework and a new impetus for evolutionary ethics. The lingering problems were the philosophical is-ought barrier and biological determinism. After tracing the early and more recent development of evolutionary ethics, I argue that the remaining problems can be surmounted and an incipient evolutionary ethics can be defended. Thoroughgoing evolutionaryethics must await theoretical developments in neurobiology and cognitive science. (shrink)

Game theory has played a critical role in elucidating the evolutionary origins of social behavior. Sober and Wilson model altruism as a prisoner's dilemma and claim that this model indicates that altruism arose from group selection pressures. Sober and Wilson also suggest that the prisoner's dilemma model can be used to characterize punishment; hence, punishment too originated from group selection pressures. However, empirical evidence suggests that a group selection model of the origins of (...) altruistic punishment may be insufficient. I argue that examining dominance hierarchies and coalition formation in chimpanzee societies suggests that the origins of altruistic punishment may be best captured by individual selection models. I suggest that this shows the necessity of coupling of game-theoretic models with a conception of what our actual social structure may have been like to best model the origins of our own behavior. ‡I would like to thank Zachary Ernst and Emma Marris for their many helpful comments which greatly improved this paper. †To contact the author, please write to: Department of Philosophy, University of Maryland, Skinner Building, College Park, MD 20742; e-mail: [email protected]. (shrink)

The development of modern evolutionary ethics began shortly after the publication of Charles Darwin's On the Origin of Species by Natural Selection. Early discussions were plagued by several problems. First, evolutionary ethical explanations were dependent on group‐selection accounts of social behavior (especially the explanation of altruism). Second, they seem to violate the philosophical principle that “ought” statements cannot be derived from “is” statements alone (values cannot be derivedfrom facts alone). Third, evolutionary ethics appeared to (...) be biologically deterministic, deemed incompatible with the free will required for ethics to be possible. Fourth, social policies based on evolutionary theory (for example, eugenics in the early part of this century) seemed patently unethical. Sociobiology (which coalesced as a field of study with Edward O. Wilson's Sociobiology: The New Synthesis, 1975) addressed several of these problems and provided a rich framework and a new impetus for evolutionary ethics. The lingering problems were the philosophical is‐ought barrier and biological determinism. After tracing the early and more recent development of evolutionary ethics, I argue that the remaining problems can be surmounted and an incipient evolutionary ethics can be defended. Thoroughgoing evolutionaryethics must await theoretical developments in neurobiology and cognitive science. (shrink)

The development of modern evolutionary ethics began shortly after the publication of Charles Darwin's On the Origin of Species by Natural Selection. Early discussions were plagued by several problems. First, evolutionary ethical explanations were dependent on group‐selection accounts of social behavior (especially the explanation of altruism). Second, they seem to violate the philosophical principle that “ought” statements cannot be derived from “is” statements alone (values cannot be derivedfrom facts alone). Third, evolutionary ethics appeared to (...) be biologically deterministic, deemed incompatible with the free will required for ethics to be possible. Fourth, social policies based on evolutionary theory (for example, eugenics in the early part of this century) seemed patently unethical. Sociobiology (which coalesced as a field of study with Edward O. Wilson's Sociobiology: The New Synthesis, 1975) addressed several of these problems and provided a rich framework and a new impetus for evolutionary ethics. The lingering problems were the philosophical is‐ought barrier and biological determinism. After tracing the early and more recent development of evolutionary ethics, I argue that the remaining problems can be surmounted and an incipient evolutionary ethics can be defended. Thoroughgoing evolutionaryethics must await theoretical developments in neurobiology and cognitive science. (shrink)

Disease represents a principal tentacle of natural selection and a staple theme of evolutionary medicine. However, it is through a small portal of entry and a very long lineage that disease as sickness entered behavioural spaces and human consciousness. This has a long evolutionary history. Anyone interested in the origins of medicine and psychiatry as social institution has to start with analysis of how mind and body were conceptualised and played out behaviourally following the pongid/hominin split and (...) thereafter. The early evolution of medicine provides a template for clarifying elemental characteristics of mind and minding. Sickness and healing in chimpanzees represents an early manifestation of (ethno) medicine, termed a behavioural tradition, which is found played out in routines of helping, caring, and healing as well as other social behaviours. Chimpanzees seem to know they are sick since they resort to self-medication when exhibiting signs and symptoms of disease. Also, they help those exhibiting physical and cognitive disability. Among hominins, awareness of consequences and implications of sickness and coping with them represented an important feature of human consciousness and a major factor in the origins of vaunted human abilities involving language, cognition, and culture as we know them. A philosophical examination of the early evolution of sickness and healing provides a window into an understanding of evolving human capacities such as self-awareness, awareness and implications of suffering, theory of mind, altruism, conceptual grasp of sickness and healing and morality. (shrink)

Anybody interested in evolutionary explanations of social phenomena (and every philosopher should be) will learn a lot from Unto Others. In addition to its cornucopia of fascinating empirical findings from biology and psychology, it is chock full of arresting perspectives, ingenious thought experiments, and clear expositions of difficult-indeed, treacherous-concepts that should be in every philosopher's kit. What philosophers will not learn, however, is the status of group selection in current evolutionary theory, because while Sober and Wilson (...) (hereafter S&W) strive intelligently to clarify and unify the issues, some of their efforts muddy the waters instead. This is mainly due to the lingering effects of ancient feuds and score-settling on both sides. So far as I can see, nobody has yet been able to come up with a judicious overview of the whole scene. Evolutionists love to pick a fight. Why propose a friendly amendment when you can claim to overthrow some major edifice of orthodoxy with your revolutionary idea? And what could be more satisfying than resurrecting a derided heresy? Stephen Jay Gould briefly tried to restore Goldschmidt's "hopeful monster" saltationism as part of his campaign for punctuated equilibrium, which was itself not as revolutionary as he claimed (Dennett, 1995); hardly a year goes by without somebody declaring that something they have discovered is, actually, a genuine instance of Lamarckianism. There is usually a grain of truth, or more, in these dramatic claims. Prions really do constitute a major exception to Crick's Central Dogma about the direction of information flow between DNA and proteins.. (shrink)

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. (shrink)

Consider the paradox of altruism: the existence of truly altruistic behaviors is difficult to reconcile with evolutionary theory if natural selection operates only on individuals, since in that case individuals should be unwilling to sacrifice their own fitness for the sake of others. Evolutionists have frequently turned to the hypothesis of group selection to explain the existence of altruism; but group selection cannot explain the evolution of morality, since morality is a one-group phenomenon (...) and group selection is a many-group phenomenon. After spelling out just what the problem is, this paper discusses several ways of solving it. (shrink)

Consider the paradox of altruism: the existence of truly altruistic behaviors is difficult to reconcile with evolutionary theory if natural selection operates only on individuals, since in that case individuals should be unwilling to sacrifice their own fitness for the sake of others. Evolutionists have frequently turned to the hypothesis of group selection to explain the existence of altruism; but group selection cannot explain the evolution of morality, since morality is a one-group phenomenon (...) and group selection is a many-group phenomenon. After spelling out just what the problem is, this paper discusses several ways of solving it. (shrink)

Discussions of altruism occur in three importantly different contexts. During the past four decades, evolutionary theory has been concerned with the possibility that forms of behaviour labelled as altruistic could emerge and could be maintained under natural selection. In these discussions, an agent A is said to act altruistically towards a beneficiary B when A's action promotes the expected reproductive success of B at expected reproductive cost to A. This sort of altruism, biological altruism, (...) is quite different from the kind of behaviour important to debates about ethical and social issues. There the focus is on psychological altruism, a notion that is concerned with the intentions of the agent and that need have no connection with the spread of anyone's genes. Psychological altruists are people with other-directed desires, emotions or intentions (this is a rough preliminary characterization, to be refined below). Finally, in certain kinds of social scientific research, the important concept is that of behavioural altruism. From the outside, behavioural altruists look like psychological altruists, although their motives and preferences may be very different. (shrink)

Advances in multiagent simulation techniques make it possible to study more realistic dynamics of complex systems and allow evolutionary theories to be tested. Here I use simulations to assess the relative importance of reproductive systems (haplodiploidy vs. diploidy), mate selection (assortative mating vs. random mating) and social economics (pay-off matrices of evolutionary games) in the evolutionary dynamics leading to the emergence of social cooperation in the provision of parental care. The simulations confirm that haplo-diploid organisms and (...) organisms mating assortatively have a higher probability for fixing alleles and require less favorable conditions for their fixation, than diploids or organisms mating randomly. The simulations showed that social behavior was most likely to emerge a) when the cost for parental investment was much lower than the benefits to the offspring, b) when cooperation improved synergistically the fitness of offspring compared to the corresponding egoistic behavior and c) when alleles coding for altruistic or social behavior could be rapidly fixed in the population, thanks to mechanisms such as haplo-diploidy and/or assortative mating. Cooperative social behavior always appeared if sociality conferred much higher fitness gains compared to non cooperative alternatives suggesting that the most important factors for the emergence and maintenance of social behavior are those based on energetic or efficiency considerations. The simulations, in congruence with the scant experimental evidence available, suggest that economic considerations rather than genetic ones are critical in explaining the emergence and maintenance of sociality. (shrink)

Humans often behave altruistically towards strangers with no chance of reciprocation. From an evolutionary perspective, this is puzzling. The evolution of altruistic cooperative behavior—in which an organism’s action reduces its fitness and increases the fitness of another organism —only makes sense when it is directed at genetically related organisms or when one can expect the favor to be returned. Therefore, evolutionary theorists such as Sober and Wilson have argued that we should revise Neo-Darwininian evolutionary theory. They (...) argue that human altruism evolved through group selection in which groups of altruists were naturally selected because they had a comparative advantage over other groups. Wilson and Sober’s hypothesis attracted followers but is rejected by most of their peers. The heated debate between advocates and critics of group selection often suffers from a lack of conceptual clarity. In response, I set out to clearly distinguish ‘genetic’ from ‘cultural’ group selection and argue that the latter does not face the potentially debilitating problems plaguing the former. I defend the claim that human altruistic dispositions evolved through cultural group selection and gene-culture coevolution and offer empirical evidence in support. I also argue that actual altruistic behavior often goes beyond the kind of behavior humans have evolved to display. Conscious and voluntary reasoning processes, I show, have an important role in altruistic behavior. This is often overlooked in the scientific literature on human altruism. (shrink)

Classical evolutionary explanations of social behavior classify behaviors from their effects, not from their underlying mechanisms. Here lies a potential objection against the view that morality can be explained by such models, e.g. Trivers’reciprocal altruism. However, evolutionary theory reveals a growing interest in the evolution of psychological mechanisms and factors them in as selective forces. This opens up perspectives for evolutionary approaches to problems that have traditionally worried moral philosophers. Once the ability to mind-read is (...) factored-in among the relevant variables in the evolution of moral abilities and counted among the selection pressures that have plausibly shaped our nature as moral agents, an evolutionary approach can contribute, so I will argue, to the solution of a long-standing debate in moral philosophy and psychology concerning the basic motivation for moral behavior. (shrink)

We propose a conceptual model that maps the causal pathways relating biological evolution to cultural change. It builds on conventional evolutionary theory by placing emphasis on the capacity of organisms to modify sources of natural selection in their environment (niche construction) and by broadening the evolutionary dynamic to incorporate ontogenetic and cultural processes. In this model, phenotypes have a much more active role in evolution than generally conceived. This sheds light on hominid evolution, on the evolution (...) of culture, and on altruism and cooperation. Culture amplifies the capacity of human beings to modify sources of natural selection in their environments to the point where that capacity raises some new questions about the processes of human adaptation. Key Words: adaptation; altruism; cooperation; evolutionary psychology; gene-culture coevolution; human evolution; human genetics; niche construction; sociobiology. (shrink)

The dominant theory of the evolution of moral cognition across a variety of fields is that moral cognition is a biological adaptation to foster social cooperation. This chapter argues, to the contrary, that moral cognition is likely an evolutionary exaptation: a form of cognition where neurobiological capacities selected for in our evolutionary history for a variety of different reasons—many unrelated to social cooperation—were put to a new, prosocial use after the fact through individual rationality, learning, and the (...) development and transmission of social norms. This chapter begins with a brief overview of the emerging behavioral neuroscience of moral cognition. It then outlines a novel theory of moral cognition that I have previously argued explains these findings better than alternatives. Finally, it shows how the evidence for this theory of moral cognition and human evolutionary history together suggest that moral cognition is likely not a biological adaptation. Instead, like reading sheet music or riding a bicycle, moral cognition is something that individuals learn to do—in this case, in response to sociocultural norms created in our ancestral history and passed down through the ages to enable cooperative living. (shrink)

The group selection controversy is about whether natural selection ever operates at the level of groups, rather than at the level of individual organisms. Traditionally, group selection has been invoked to explain the existence of altruistic behaviour in nature. However, most contemporary evolutionary biologists are highly sceptical of the hypothesis of group selection, which they regard as biologically implausible and not needed to explain the evolution of altruism anyway. But in their recent book, Elliot (...) Sober and David Sloan Wilson [1998] argue that the widespread opposition to group selection is founded on conceptual confusion. The theories that have been propounded as alternatives to group selection are actually group selection in disguise, they maintain. I examine their arguments for this claim, and John Maynard Smith's arguments against it. I argue that Sober and Wilson arrive at a correct position by faulty reasoning. In the final section, I examine the issue of how to apply the principle of natural selection at different levels of the biological hierarchy, which underlies the dispute between Sober and Wilson and Maynard Smith. (shrink)

The authors demonstrate that unselfish behavior is in fact an important feature of both biological and human nature. Their book provides a panoramic view of altruism throughout the animal kingdom--from self-sacrificing parasites to the human capacity for selflessness--even as it explains the evolutionary sense of such behavior.

While recent developments in evolutionary theory, particularly game-theoretical models of group selection, can appear to provide a potential evolutionary grounding for human altruism, significant ethical problems remain embedded in such portrayals of human interaction. Specifically, such models end up treating the value of the individual as subservient to group survival, rather than viewing each unique individual as an ‘end in herself’. As such, a contradiction remains between the picture of human relationships that arises from (...) class='Hi'>evolutionary game-theoretical accounts and the picture that arises from many ethical and religious accounts, particularly those linked to the biblical conception of the image of God. However, I argue that such contradictions between scientific and ethical-religious pictures need not lead to despair; rather, by drawing a parallel to dialogues between individuals holding ‘contradictory’ theological claims, we can discern ways in which such contradiction can generate a fruitful tension that can lead to deeper understanding both of one’s own tradition and that of the other person. Accordingly, as with inter-faith dialogue, interactions between evolutionary theorists and religious ethicists can also give rise to greater self-understanding and expanded intellectual horizons, not in spite of but precisely by virtue of their ‘incompatible’ pictures of the human being. (shrink)

Inclusive fitness theory provides conditions for the evolutionary success of a gene. These conditions ensure that the gene is selfish in the sense of Dawkins (The selfish gene, Oxford University Press, Oxford, 1976): genes do not and cannot sacrifice their own fitness on behalf of the reproductive population. Therefore, while natural selection explains the appearance of design in the living world (Dawkins in The blind watchmaker: why the evidence of evolution reveals a universe without design, W. W. (...) Norton, New York, 1996), inclusive fitness theory does not explain how. Indeed, Hamilton’s rule is equally compatible with the evolutionary success of prosocial altruistic genes and antisocial predatory genes, whereas only the former, which account for the appearance of design, predominate in successful organisms. Inclusive fitness theory, however, permits a formulation of the central problem of sociobiology in a particularly poignant form: how do interactions among loci induce utterly selfish genes to collaborate, or to predispose their carriers to collaborate, in promoting the fitness of their carriers? Inclusive fitness theory, because it abstracts from synergistic interactions among loci, does not answer this question. Fitness-enhancing collaboration among loci in the genome of a reproductive population requires suppressing alleles that decrease, and promoting alleles that increase the fitness of its carriers. Suppression and promotion are effected by regulatory networks of genes, each of which is itself utterly selfish. This implies that genes, and a fortiori individuals in a social species, do not maximize inclusive fitness but rather interact strategically in complex ways. It is the task of sociobiology to model these complex interactions. (shrink)

Evolutionary applications of game theory present one of the most pedagogically accessible varieties of genuine, contemporary theoretical biology. We present here Oyun (OY-oon, http://charlespence.net/oyun), a program designed to run iterated prisoner’s dilemma tournaments, competitions between prisoner’s dilemma strategies developed by the students themselves. Using this software, students are able to readily design and tweak their own strategies, and to see how they fare both in round-robin tournaments and in “evolutionary” tournaments, where the scores in a given “generation” (...) directly determine contribution to the population in the next generation. Oyun is freely available, runs on Windows, Mac, and Linux computers, and the process of creating new prisoner’s dilemma strategies is both easy to teach and easy for students to grasp. We illustrate with two interesting examples taken from actual use of Oyun in the classroom. (shrink)

Species evolve, very slowly, through selection of genes which give rise to phenotypes well adapted to their environments. The cultures, including the languages, of human communities evolve, much faster, maintaining at least a minimum level of adaptedness to the external, non- cultural environment. In the phylogenetic evolution of species, the transmission of information across generations is via copying of molecules, and innovation is by mutation and sexual recombination. In cultural evolution, the transmission of information across generations is by learning, (...) and innovation is by sporadic invention or borrowing from other cultures. This much is the foundational bedrock of evolutionary theory. But things get more complicated; there can be gene-culture co-evolution. Prior to the rise of culture, the physical environment is the only force shaping biological evolution from outside the organism, and cultures themselves are clearly constrained by the evolved biological characteristics of their members. But cultures become part of the external environment, and influence the course of biological evolution. For example, altruistic cultures with developed medical knowledge reduce the cost to the individual of carrying genes disposing to certain pathologies (such as diabetes); and such genes become more widespread in the populations maintaining such cultures. Assortative mating can affect biological evolution, and particular cultures may influence the factors which are sorted for in mating. (For a careful discussion of the effects of cultural evolution on natural selection, see Cavalli-Sforza and Bodmer, 1971:774- 804). This paper examines mechanisms involved in the co-evolution of a biological trait, the critical period for language acquisition, and a property of human cultures, the size of their languages. A gene/culture interaction will be shown that can be described as a kind of symbiosis, but perhaps more aptly as an `arms race'. In this introduction, we will sketch the basic mechanics of the interaction in very broad terms; the rest of the paper will explain and justify the details. The implications of our model for second language acquisition are given toward the end of the paper. (shrink)

The use of evolutionary game theory to explain the evolution of human norms and the behavior of humans who act according to those norms is widespread. Both the aims and motivation for its use are clearly articulated by Harms and Skyrms (2008) in the following passage: "A good theory of evolution of norms might start by explaining the evolution of altruism in Prisoner’s Dilemma, of Stag Hunting, and of the equal split in the symmetric bargaining game. (...) These are not well-explained by classical game theory based on rational choice. From a technical point of view, they present different theoretical challenges. In the bargaining game, there are an infinite number of equilibria with no principled (rational choice) way to select the cooperative one. In Stag Hunt there are only two, but the non-cooperative one is selected by risk-dominance. In Prisoner’s Dilemma the state of mutual cooperation is not a Nash equilibrium at all, and cooperation flies in the face of the rational-choice principle that one does not choose less rather than more. In contrast to rational choice theory, the most common tool of evolutionary game theory is the replicator dynamics, in which the propagation rate of each strategy is determined by its current payoffs. These dynamics have a rationale in both biological and cultural evolutionary modeling, and sometimes tell us things that rational choice theory does not." We agree with the first sentence in this quotation: a good theory about the behavior under norms ought to explain altruism in the Prisoner’s Dilemma (PD), playing Stag in Stag Hunt (SH), and offering equal splits in the symmetric Nash bargaining game (NB). We also agree with Harms and Skyrms about the difference in technical challenges each of these games poses. Finding a single mechanism, even one as broadly understood as evolution, that could solve these challenges en masse is no doubt a tall order. Nonetheless, in this paper, we present a single, simple, modification to SH, NB, and a general n-player PD that does just that: we introduce deontological autonomy into the models. (shrink)

Modern scientific theories of emotional behavior, almost without exception, trace their origin to Charles Darwin, and his publications On the Origin of Species (1859) and The Expression of the Emotions in Man and Animals (1872). The most famous evolutionary dilemma Darwin acknowledged as a challenge to his theory of natural selection was the incomplete sub Cambrian fossil record. However, Darwin struggled with two other rarely referenced theoretical and scientific dilemmas that confounded his theories about emotional behavior. These (...) included (1) the origin of social instincts (e.g., altruism, empathy, reciprocity and cooperation) and the reasons for their conservation in evolution and (2) the peripheral control of heart rate vis-a-vis emotional behavior outside of consciousness. Darwin acknowledged that social instincts are critical to the survival of some species, but had difficulty aligning them with his theory of natural selection in humans. Darwin eventually proposed that heart rate and emotions are controlled via one’s intellect and cortical mechanisms, and that instinctive behavior is genetically-programmed and inherited. Despite ongoing efforts, these two theoretical dilemmas are debated to this day. Simple testable hypotheses have yet to emerge for the biological mechanisms underlying instinctive behavior or the way heart rate is controlled in infants. In this paper, we review attempts to resolve ¬these issues over the past 160 years. We posit that research and theories that supported Darwin’s individualistic brain-centric and genetic model have become an “orthodox” Western view of emotional behavior, one that produced the prevailing behavioral construct of attachment as developed by John Bowlby. We trace research and theories that challenged this orthodoxy at various times, and show how these challenges were repeatedly overlooked, rejected, or misinterpreted. We review two new testable theories, emotional connection theory and calming cycle theory, which we argue resolve the two dilemmas We show emerging scientific evidence from physiology and a wide variety of other fields, as well from clinical trials among prematurely born infants, that supports the two theories. Clinical implications of the new theories and possible new ways to assess risk and intervene in emotional, behavioral and developmental disorders are discussed. (shrink)

The eminent historian and philosopher of biology, Michael Ruse, has written several books that explore the relationship of evolutionary theory to its larger scientific and cultural setting. Among the questions he has investigated are: Is evolution progressive? What is its epistemological status? Most recently, in "Darwin and Design: Does Evolution have a Purpose?," Ruse has provided a history of the concept of teleology in biological thinking, especially in evolutionary theorizing. In his book, he moves quickly from Plato (...) and Aristotle to Kant and such British thinkers as Paley and Whewell. His main focus, though, is on Darwin's theory and its subsequent fate. Ruse rests his history on some shaky historical and philosophic assumptions, particularly the unexamined notion that evolutionary theory is an abstract entity that is unproblematically realized in different historical periods. He also assumes that Darwin conceived nature as if it were a Manchester spinning loom -- a clanking, dispassionate machine. A more subtle analysis, which Ruse eschews, might discover that Darwin's conception of nature owed a strong debt to German Romanticism and that he contrived to infuse nature with moral and aesthetic values, not to suck them from nature. Ruse proves he is a thinker to contend with, and this essay is quite contentious. (shrink)

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. (shrink)

Although moral psychologists and feminist moral theorists emphasize males’ interest in justice or fairness and females’ interest in care or empathy, recent work in evolutionary psychology links females’ interests in care and empathy for others with interests in fairness and equality. In an important work on sex differences in cognitive abilities, David Geary (1998) argues that the evolutionary mechanism of sexual selection drives the evolution of particular cognitive abilities and selection for particular interests. I mount two (...) main challenges to Geary’s claims. First, I argue that female social and intrasexual competitive environments evolve, which challenges the assumption that such environments are largely nonkin-based and characterized by reciprocity. Second, I grant Geary’s entire characterization of female environments, but argue that the natures of reciprocal relationships themselves do not require and may not select for interests in fairness and equality. This analysis not only challenges claims regarding sex differences in moral interests, but also suggests the need for a diachronic model of male and female social and intrasexual competitive environments. In addition, I propose a return to Trivers’s (1971) focus on the suite of emotions underlying reciprocal altruism as properties and features of individuals as fodder for selection. (shrink)

Many of the most important properties of human groups – including properties that may give one group an evolutionary advantage over another – are properly defined only at the level of group organization. Yet at present, most work on the evolution of culture has focused solely on the transmission of individual-level traits. I propose a conceptual extension of the theory of cultural evolution, particularly related to the evolutionary competition between cultural groups. The key concept in this extension (...) is the emergent group-level trait. This type of trait is characterized by the structured organization of differentiated individuals and constitutes a unit of selection that is qualitatively different from selection on groups as defined by traditional multilevel selection theory. As a corollary, I argue that the traditional focus on cooperation as the defining feature of human societies has missed an essential feature of cooperative groups. Traditional models of cooperation assume that interacting with one cooperator is equivalent to interacting with any other. However, human groups involve differential roles, meaning that receiving aid from one individual is often preferred to receiving aid from another. In this target article, I discuss the emergence and evolution of group-level traits and the implications for the theory of cultural evolution, including ramifications for the evolution of human cooperation, technology, and cultural institutions, and for the equivalency of multilevel selection and inclusive fitness approaches. (shrink)

It is a challenge to explain how evolutionary altruism can evolve by the process of natural selection, since altruists in a group will be less fit than the selfish individuals in the same group who receive benefits but do not make donations of their own. Darwin proposed a theory of group selection to solve this puzzle. Very simply, even though altruists are less fit than selfish individuals within any single group, groups of altruists are more (...) fit than groups of selfish individuals. If a population is subdivided into many groups that vary in their altruistic tendencies, altruism will be favored at the level of selection among groups even as it is being disfavored at the level of selection among individuals within groups. Darwin’s scenario became the basis for a theoretical framework called multilevel selection theory. (shrink)

It is a challenge to explain how evolutionary altruism can evolve by the process of natural selection, since altruists in a group will be less fit than the selfish individuals in the same group who receive benefits but do not make donations of their own. Darwin proposed a theory of group selection to solve this puzzle. Very simply, even though altruists are less fit than selfish individuals within any single group, groups of altruists are more (...) fit than groups of selfish individuals. If a population is subdivided into many groups that vary in their altruistic tendencies, altruism will be favored at the level of selection among groups even as it is being disfavored at the level of selection among individuals within groups. Darwin’s scenario became the basis for a theoretical framework called multilevel selection theory. (shrink)

It is a challenge to explain how evolutionary altruism can evolve by the process of natural selection, since altruists in a group will be less fit than the selfish individuals in the same group who receive benefits but do not make donations of their own. Darwin proposed a theory of group selection to solve this puzzle. Very simply, even though altruists are less fit than selfish individuals within any single group, groups of altruists are more (...) fit than groups of selfish individuals. If a population is subdivided into many groups that vary in their altruistic tendencies, altruism will be favored at the level of selection among groups even as it is being disfavored at the level of selection among individuals within groups. Darwin’s scenario became the basis for a theoretical framework called multilevel selection theory. (shrink)