Questioning the cultural evolution of altruism

The evolutionary foundations of helping among nonkin in humans have been the object of intense debates in the past decades. One thesis has had a prominent influence in this debate: the suggestion that genuine altruism, strictly defined as a form of help that comes at a net fitness cost for the benefactor, might have evolved owing to cultural transmission. The gene–culture coevolution literature is wont to claim that cultural evolution changes the selective pressures that normally act to limit the emergence of altruistic behaviours. This paper aims to recall, however, that cultural transmission yields altruism only to the extent that it relies on maladaptive mechanisms, such as conformist imitation and (in some cases) payoff‐biased transmission. This point is sometimes obscured in the literature by a confusion between genuine altruism, maladaptive by definition, and mutualistic forms of cooperation, that benefit all parties in the long run. Theories of cultural altruism do not lift the selective pressures weighing on strictly altruistic actions; they merely shift the burden of maladaptation from social cognition to cultural transmission.     

The Significance of Non-vertical Transmission of Phenotype for the Evolution of Altruism

My aim in this paper is to demonstrate that a very simple learning rule based on imitation can help to sustain altruism as a culturally transmitted pattern or behaviour among agents playing a standard prisoner’s dilemma game. The point of this demonstration is not to prove that imitation is single-handedly responsible for existing levels of altruism (a thesis that is false), nor is the point to show that imitation is an important factor in explanations for the evolution of altruism (a thesis already prominent in the existing literature). The point is to show that imitation contributes to the evolution of altruism in a particular way that is not always fairly represented by evolutionary game theory models. Specifically, the paper uses a simple model to illustrate that cultural transmission includes mechanisms that do not transmit phenotype vertically (i.e. from parent to related offspring) and that these mechanisms can promote altruism in the absence of any direct biological propensity favouring such behaviour. This is a noteworthy result because it shows that evolutionary models can be built to explicitly reflect the contribution of non-vertical transmission in our explanations for the evolution of altruism among humans and other social species.     

Altruism across disciplines: one word, multiple meanings

Altruism is a deep and complex phenomenon that is analysed by scholars of various disciplines, including psychology, philosophy, biology, evolutionary anthropology and experimental economics. Much confusion arises in current literature because the term altruism covers variable concepts and processes across disciplines. Here we investigate the sense given to altruism when used in different fields and argumentative contexts. We argue that four distinct but related concepts need to be distinguished: (a) psychological altruism, the genuine motivation to improve others’ interests and welfare; (b) reproductive altruism, which involves increasing others’ chances of survival and reproduction at the actor’s expense; (c) behavioural altruism, which involves bearing some cost in the interest of others; and (d) preference altruism, which is a preference for others’ interests. We show how this conceptual clarification permits the identification of overstated claims that stem from an imprecise use of terminology. Distinguishing these four types of altruism will help to solve rhetorical conflicts that currently undermine the interdisciplinary debate about human altruism.     

Sacrifice, Plants, and Western Pharmaceuticals: Money and Health Care in Northern Ghana

In this article I discuss the role of money vis-à-vis health care among the Dagomba, an agrarian people living in northern Ghana, whose pluralistic medical culture involves the use of both plants and Western pharmaceuticals in the treatment of various symptoms. In Dagomba society monetary exchanges in the domain of healing cannot be equated with self-interest, and nonmonetary exchanges cannot be compared with altruism in any straightforward fashion. Exchanges and their purposes are made meaningful by the contexts in which they occur. Exchanges may involve money and be commoditized. However, the very fact that such an exchange is made in a "benevolent and considerate manner" (which often means selling/buying on credit) transforms otherwise impersonal relationships between medicine sellers and their customers into displays of solidarity with the communal (collective) goals of Dagomba society.     

Altruism and Reward: Motivational Compatibility in Deceased Organ Donation

Acts of helping others are often based on mixed motivations. Based on this claim, it has been argued that the use of a financial reward to incentivize organ donation is compatible with promoting altruism in organ donation. In its report Human Bodies: Donation for Medicine and Research, the Nuffield Council on Bioethics uses this argument to justify its suggestion to pilot a funeral payment scheme to incentivize people to register for deceased organ donation in the UK. In this article, I cast a sceptical eye on the above Nuffield report's argument that its proposed funeral payment scheme would prompt deceased organ donations that remain altruistic (as defined by and valued the report). Specifically, I illustrate how this scheme may prompt various forms of mixed motivations which would not satisfy the report's definition of altruism. Insofar as the scheme produces an expectation of the reward, it stands diametrical to promoting an ‘altruistic perspective’. My minimal goal in this article is to argue that altruism is not motivationally compatible with reward as an incentive for donation. My broader goal is to argue that if a financial reward is used to incentivize organ donation, then we should recognize that the donation system is no longer aiming to promote altruism. Rewarded donation would not be altruistic but it may be ethical given a persistent organ shortage situation.     

Multilevel selection, cooperation, and altruism

Unto Others (Sober and Wilson 1998) shows how the general principles of Multi-Level Selection (MLS) theory apply to selection at multiple levels of the biological hierarchy. It also argues for the existence of "genuine" evolutionary and psychological altruism. The authors’ views on altruism do not follow logically from principles of MLS, and their failure do disentangle these two themes undermines their otherwise excellent presentation of MLS theory. Rebuttal of the view that human nature is completely selfish depends not on the prevalence of altruism but on the importance of group-advantageous traits that benefit both self and other group members without necessarily inflicting direct costs on outsiders.     

On the relationship between evolutionary and psychological definitions of altruism and selfishness

I examine the relationship between evolutionary definitions of altruism that are based on fitness effects and psychological definitions that are based on the motives of the actor. I show that evolutionary altruism can be motivated by proximate mechanisms that are psychologically either altruistic or selfish. I also show that evolutionary definitions do rely upon motives as a metaphor in which the outcome of natural selection is compared to the decisions of a psychologically selfish (or altruistic) individual. Ignoring the precise nature of both psychological and evolutionary definitions has obscured many important issues, including the biological roots of psychological altruism.     

The conjoined twins and the limits of rationality in applied ethics

In this article I consider the case of the surgical separation of conjoined twins resulting in the immediate and predictable death of the weaker one. The case was submitted to English law by the hospital, and the operation permitted against the parents' wishes. I consider the relationship between the legal decision and the moral reasons adduced in its support, reasons gaining their force against the framework of much mainstream normative ethical theory. I argue that in a few morally dilemmatic situations, such a legalistic–theoretical approach cannot plausibly accommodate certain irreducible and ineliminable features of the ethical experience of any concrete individual implicated in the situation, and that this failure partly undermines its self–appointed role of guiding such an individual's conduct. For example, the problem as experienced by the judge and by the parents might not be the same problem at all, and some of their respective reasons may be mutually unintelligible or impotent. I certainly do not argue for a rejection of law or of moral theory; I merely challenge their implicit claim to comprehensiveness and their fixation with an idealised and putatively universal rationality modelled on converging scientific enquiry. Finally, I claim that at least in the twins' case there may be insufficient normative robustness to the conclusions reached, or indeed reachable, by the court in a situation where intuitions and moral reasons pull in fundamentally incommensurable directions; as such, there may be room for an acknowledgement of the spiritual, through a humble abstention from making a decision – which is not to be confused with deciding to do nothing.     

Sex-biased dispersal, haplodiploidy and the evolution of helping in social insects

In his famous haplodiploidy hypothesis, W. D. Hamilton proposed that high sister-sister relatedness facilitates the evolution of kin-selected reproductive altruism among Hymenopteran females. Subsequent analyses, however, suggested that haplodiploidy cannot promote altruism unless altruists capitalize on relatedness asymmetries by helping to raise offspring whose sex ratio is more female-biased than the population at large. Here, we show that haplodiploidy is in fact more favourable than is diploidy to the evolution of reproductive altruism on the part of females, provided only that dispersal is male-biased (no sex-ratio bias or active kin discrimination is required). The effect is strong, and applies to the evolution both of sterile female helpers and of helping among breeding females. Moreover, a review of existing data suggests that female philopatry and non-local mating are widespread among nest-building Hymenoptera. We thus conclude that Hamilton was correct in his claim that 'family relationships in the Hymenoptera are potentially very favourable to the evolution of reproductive altruism'.     

Why reciprocal altruism is not a kind of group selection

Reciprocal altruism was originally formulated in terms of individual selection and most theorists continue to view it in this way. However, this interpretation of reciprocal altruism has been challenged by Sober and Wilson (1998). They argue that reciprocal altruism (as well as all other forms of altruism) evolves by the process of group selection. In this paper, we argue that the original interpretation of reciprocal altruism is the correct one. We accomplish this by arguing that if fitness attaches to (at minimum) entire life cycles, then the kind of fitness exchanges needed to form the group-level in such situations is not available. Reciprocal altruism is thus a result of individual selection and when it evolves, it does so because it is individually advantageous.     

Mistakes allow evolutionary stability in the repeated prisoner's dilemma game

The repeated prisoner's dilemma game has been widely used in analyses of the evolution of reciprocal altruism. Recently it was shown that no pure strategy could be evolutionarily stable in the repeated prisoner's dilemma. Here I show that if there is always some probability that individuals will make a mistake, then a pure strategy can be evolutionarily stable provided that it is "strong perfect equilibria" against itself. To be a strong perfect equilibrium against itself, a strategy must be the best response to itself after every possible sequence of behavior. I show that both unconditional defection and a modified version of tit-for-tat have this property.     

Pathogen burden,co‐infection and major histocompatibility complex variability in the European badger (Meles meles)

Pathogen‐mediated selection is thought to maintain the extreme diversity in the major histocompatibility complex (MHC) genes, operating through the heterozygote advantage, rare‐allele advantage and fluctuating selection mechanisms. Heterozygote advantage (i.e. recognizing and binding a wider range of antigens than homozygotes) is expected to be more detectable when multiple pathogens are considered simultaneously. Here, we test whether MHC diversity in a wild population of European badgers (Meles meles) is driven by pathogen‐mediated selection. We examined individual prevalence (infected or not), infection intensity and co‐infection of 13 pathogens from a range of taxa and examined their relationships with MHC class I and class II variability. This population has a variable, but relatively low, number of MHC alleles and is infected by a variety of naturally occurring pathogens, making it very suitable for the investigation of MHC–pathogen relationships. We found associations between pathogen infections and specific MHC haplotypes and alleles. Co‐infection status was not correlated with MHC heterozygosity, but there was evidence of heterozygote advantage against individual pathogen infections. This suggests that rare‐allele advantages and/or fluctuating selection, and heterozygote advantage are probably the selective forces shaping MHC diversity in this species. We show stronger evidence for MHC associations with infection intensity than for prevalence and conclude that examining both pathogen prevalence and infection intensity is important. Moreover, examination of a large number and diversity of pathogens, and both MHC class I and II genes (which have different functions), provide an improved understanding of the mechanisms driving MHC diversity.     

Consequentialism, reasons, value and justice

Over the past 10 years, John Harris has made important contributions to thinking about distributive justice in health care. In his latest work, Harris controversially argues that clinicians should stop prioritising patients according to prognosis. He argues that the good or benefit of health care is providing each individual with an opportunity to live the best and longest life possible for him or her. I call this thesis, opportunism. For the purpose of distribution of resources in health care, Harris rejects welfarism (the thesis that the good of health care is well-being) and argues that utilitarianism in general may lead to de facto discrimination against groups of people needing health care. I argue that well-being is a superior theory of the good of health care to Harris' opportunism. Harris' concerns about utilitarianism can be better addressed by: (i) relating justice more closely to reasons for action; (ii) by conceptualising the relationship between reasons for action and the value of the consequences of those actions as a plateau rather than scalar relationship. Justice can be understood as satisfying as many equally rational claims on resources as possible. The rationality of a person's claim on health resources turns on the strength of that person's reasons to promote certain health-related states of affairs. I argue that the strength of that reason does not track the expected value of that state of affairs in a fully scalar fashion. Rather a person can have most reason to promote some state of affairs, even though he or she could promote other more valuable states of affairs. Thus there can be equal reason for a distributor of public resources to save either of two people, even though one will have a better and more valuable life. This approach, while addressing many of Harris' concerns about utilitarianism, does not imply that doctors should give up prioritising patients according to prognosis altogether, but it does allow that patients with lower but reasonable prognosis should have a share of public resources.     

Wrongs, preferences, and the selection of children: a critique of Rebecca Bennett's argument against the principle of procreative beneficence

Rebecca Bennett, in a recent paper dismissing Julian Savulescu's principle of procreative beneficence, advances both a negative and a positive thesis. The negative thesis holds that the principle's theoretical foundation - the notion of impersonal harm or non-person-affecting wrong - is indefensible. Therefore, there can be no obligations of the sort that the principle asserts. The positive thesis, on the other hand, attempts to plug an explanatory gap that arises once the principle has been rejected. That is, it holds that the intuitions of those who adhere to the principle are not genuine moral intuitions, but instead simply give voice to mere (non-moral) preferences. This paper, while agreeing that Savulescu's principle does not express a genuine moral obligation, takes issue with both of Bennett's theses. It is suggested that the argument for the negative thesis is either weak or question-begging, while there is insufficient reason to suppose the positive thesis true.     

Scientific progress and the prospects for culture-bound syndromes

This paper aims to show that the classification by the American Psychiatric Association (APA) in the Diagnostic and Statistical Manual of Mental Disorders (DSM) of a distinct listing of disorders known as Culture-Bound Syndromes (CBS) is misguided. I argue that the list of CBS (in Appendix I of the manual) comprises either (a) genuine disorders that should be included within the main body of the DSM; or (b) ersatz-disorders that serve a practical role for psychiatrists dealing with patients from certain cultures but will one day be eliminated or assimilated by bona fide DSM classifications. In support of these views I draw on claims from two key themes in the philosophy of science: (1) the claim that all folk (that is, non-scientific) explanations for phenomena are thoroughly theoretical and therefore fallible; and (2) the occurrence of theoretical elimination in the history of science. I contend that any ersatz-disorders located in the DSM that are judged to be radically false do not differ in kind from eliminated theories in the history of pre-science.     

Competitive altruism: from reciprocity to the handicap principle

Current work on cooperation is focused on the theory of reciprocal altruism. However, reciprocity is just one way of getting a return on an investment in altruism and is difficult to apply to many examples. Reciprocity theory addresses how animals respond dynamically to others so as to cooperate without being exploited. I discuss how introducing differences in individual generosity together with partner choice into models of reciprocity can lead to an escalation in altruistic behaviour. Individuals may compete for the most altruistic partners and non-altruists may become ostracized. I refer to this phenomenon as competitive altruism and propose that it can represent a move away from the dynamic responsiveness of reciprocity. Altruism may be rewarded in kind, but rewards may be indirectly accrued or may not involve the return of altruism at all, for example if altruists tend to be chosen as mates. This variety makes the idea of competitive altruism relevant to behaviours which cannot be explained by reciprocity. I consider whether altruism might act as a signal of quality, as proposed by the handicap principle. I suggest that altruistic acts could make particularly effective signals because of the inherent benefits to receivers. I consider how reciprocity and competitive altruism are related and how they may be distinguished.     

The Russo–Williamson Theses in the social sciences: Causal inference drawing on two types of evidence

This article examines two theses formulated by Russo and Williamson (2007) in their study of causal inference in the health sciences. The two theses are assessed against evidence from a specific case in the social sciences, i.e., research on the institutional determinants of the aggregate unemployment rate. The first Russo–Williamson Thesis is that a causal claim can only be established when it is jointly supported by difference-making and mechanistic evidence. This thesis is shown not to hold. While researchers in my case study draw extensively on both types of evidence, one causal claim out of the three analyzed is established even though it is exclusively supported by mechanistic evidence. The second Russo–Williamson Thesis is that standard accounts of causality fail to handle the dualist epistemology highlighted in the first Thesis. I argue that a counterfactual-manipulationist account of causality—which is endorsed by many philosophers as well as many social scientists—can perfectly make sense of the typical strategy in my case study to draw on both difference-making and mechanistic evidence; it is just an instance of the common strategy of increasing evidential variety.     

Evolution of eusociality by kin selection: the effect of inbreeding between siblings

The effect of sib-sib inbreeding on the evolution of eusocial altruism in Hymenoptera by kin selection is examined by computer simulations. Inbreeding has minor effects on the ratio of relatedness to siblings: relatedness to offspring, but this ratio remains approximately one no matter what the degree of inbreeding. This implies that although inbreeding increases relatedness to siblings, relatedness to offspring increases to the same degree. Hence, inbreeding does not make the evolution of altruism more likely. If all the brothers of (non-mating) altruists outbreed, thereby increasing the frequency of altruism alleles in the outbred fraction of the population especially at low gene frequency, then altruism can be promoted by inbreeding. However, this is an indirect advantage, not attributable to inbreeding per se.     

Altruism Can Be Assessed Correctly Based on Impression

Detection of genuine altruists could be a solution to the problem of subtle cheating. Brown et al. (Evol Psychol 1:42–69, 2003) found that humans could detect altruists using nonverbal cues. However, their experiments can be improved upon in several ways, and further investigation is needed to determine whether altruist-detection abilities are human universals. In our experiment, we used video clips of natural conversations as the stimulus. We asked a sample of Japanese undergraduates to rate their own level of altruism and then to estimate the videotaped targets’ altruism using the same scale. The perceivers were able to estimate the targets’ altruism levels accurately. Perceivers’ altruism score did not affect their ability to discriminate between altruists and non-altruists. Perceivers’ impressions of the altruist and non-altruist targets were also found to be different. Coding of nonverbal behavior of the targets revealed that altruists exhibited more “felt smiles” than non-altruists, which also supports the results of the previous study.