The natural selection of altruistic traits

Proponents of the standard evolutionary biology paradigm explain human “altruism” in terms of either nepotism or strict reciprocity. On that basis our underlying nature is reduced to a function of inclusive fitness: human nature has to be totally selfish or nepotistic. Proposed here are three possible paths to giving costly aid to nonrelatives, paths that are controversial because they involve assumed pleiotropic effects or group selection. One path is pleiotropic subsidies that help to extend nepotistic helping behavior from close family to nonrelatives. Another is “warfare”—if and only if warfare recurred in the Paleolithic. The third and most plausible hypothesis is based on the morally based egalitarian syndrome of prehistoric hunter-gatherers, which reduced phenotypic variation at the within-group level, increased it at the between-group level, and drastically curtailed the advantages of free riders. In an analysis consistent with the fundamental tenets of evolutionary biology, these three paths are evaluated as explanations for the evolutionary development of a rather complicated human social nature. This paper (in a series of drafts) has profited from comments by Michael Boehm, Donald T. Campbell, Bruce Knauft, Jane Lancaster, Martin Muller, Peter J. Richerson, Gary Seaman, Craig Stanford, George Williams, Edward O. Wilson, David Sloan Wilson, and two reviewers for Human Nature. Christopher Boehm is a professor of anthropology and the director of the Jane Goodall Research Center, University of Southern California. His research interests in political anthropology concern egalitarianism, feuding, warfare, and conflict resolution (humans and chimpanzees). In biosocial anthropology he is interested in altruism, group selection, and decisions.     

The concept of group heritability

This paper investigates the role of the concept of group heritability in group selection theory, in relation to the well-known distinction between type 1 and type 2 group selection (GS1 and GS2). I argue that group heritability is required for the operation of GS1 but not GS2, despite what a number of authors have claimed. I offer a numerical example of the evolution of altruism in a multi-group population which demonstrates that a group heritability coefficient of zero is perfectly compatible with the successful operation of group selection in the GS2 sense. A diagnosis of why group heritability has wrongly been regarded as necessary for GS2 is suggested.     

Virulence and transmission modes in metapopulations: when group selection increases virulence

Individuals that are infected by a pathogen can transmit it to unrelated conspecifics (horizontal transmission) or to their progeny when they reproduce (vertical transmission). The mechanisms of these two routes of transmission are different and this difference impacts the way virulence evolves in pathogens. More precisely, horizontal transmission depends on the probability that an infected host contacts susceptible conspecifics, and therefore on its lifespan. Vertical transmission additionally depends on the host's fecundity. This additional dependence in vertically transmitted pathogens results in a decrease in their evolutionarily stable (ES) virulence.Spatial structure is another factor that is often supposed to decrease pathogens’ ES virulence, mostly because it impedes competition for transmission in local populations of hosts. In this paper, using the adaptive dynamics framework, we show that spatial structure can increase ES virulence when pathogens are mostly vertically transmitted. This is due to the difference in how pathogens compete for transmission in local population of hosts, depending on how they are transmitted. We also show that symbionts that are horizontally transmitted should respond more to a change in spatial structure than symbionts that are vertically transmitted.     

The history of Hayek’s theory of cultural evolution

This paper traces the historical origins of Friedrich A. Hayek’s theory of cultural evolution, and argues that Hayek’s evolutionary thought was significantly inspired by Alexander M. Carr-Saunders and Oxford zoology. While traditional Hayek scholarship emphasizes the influence of Carl Menger and the British eighteenth-century moral philosophers, I claim that these sources underdetermine what was most characteristic of Hayek’s theory, viz. the idea that cultural evolution is a matter of group selection, and the idea that natural selection operates on acquired as well as on inherited properties.     

Why kin and group selection models may not be enough to explain human other-regarding behaviour

Models of kin or group selection usually feature only one possible fitness transfer. The phenotypes are either to make this transfer or not to make it and for any given fitness transfer, Hamilton's rule predicts which of the two phenotypes will spread. In this article we allow for the possibility that different individuals or different generations face similar, but not necessarily identical possibilities for fitness transfers. In this setting, phenotypes are preference relations, which concisely specify behaviour for a range of possible fitness transfers (rather than being a specification for only one particular situation an animal or human can be in). For this more general set-up, we find that only preference relations that are linear in fitnesses can be explained using models of kin selection and that the same applies to a large class of group selection models. This provides a new implication of hierarchical selection models that could in principle falsify them, even if relatedness--or a parameter for assortativeness--is unknown. The empirical evidence for humans suggests that hierarchical selection models alone are not enough to explain their other-regarding or altruistic behaviour.     

A new group-selection model for the evolution of homosexuality

Abstract. Scientists have long puzzled over how homosexual orientation has evolved, given the assumed low relative fitness of homosexual individuals compared to heterosexual individuals. A number of theoretical models for the evolution of homosexuality have been postulated including balance polymorphism, "Fertile females", hypervariability of DNA sequences, kin selection, and "parental manipulation". In this paper, I propose a new group-selection model for the evolution of homosexuality which offers two advantages over existing models: (1) its non-assumption of genetic determinism, and (2) its lack of dependency on an inefficient altruism relation and family dynamics theory.     

Evolution of cooperation and altruistic punishment when retaliation is possible

Altruistic punishment is suggested to explain observed high levels of cooperation among non-kin related humans. However, laboratory experiments as well as ethnographic evidence suggest that people might retaliate if being punished, and that this reduces the level of cooperation. Building on existing models on the evolution of cooperation and altruistic punishment, we explore the consequences of the option of retaliation. We find that cooperation and altruistic punishment does not evolve with larger population levels if the option of retaliation is included.     

Rates of cultural change and patterns of cultural accumulation in stochastic models of social transmission

Cultural variation in a population is affected by the rate of occurrence of cultural innovations, whether such innovations are preferred or eschewed, how they are transmitted between individuals in the population, and the size of the population. An innovation, such as a modification in an attribute of a handaxe, may be lost or may become a property of all handaxes, which we call “fixation of the innovation.” Alternatively, several innovations may attain appreciable frequencies, in which case properties of the frequency distribution—for example, of handaxe measurements—is important. Here we apply the Moran model from the stochastic theory of population genetics to study the evolution of cultural innovations. We obtain the probability that an initially rare innovation becomes fixed, and the expected time this takes. When variation in cultural traits is due to recurrent innovation, copy error, and sampling from generation to generation, we describe properties of this variation, such as the level of heterogeneity expected in the population. For all of these, we determine the effect of the mode of social transmission: conformist, where there is a tendency for each naïve newborn to copy the most popular variant; pro-novelty bias, where the newborn prefers a specific variant if it exists among those it samples; one-to-many transmission, where the variant one individual carries is copied by all newborns while that individual remains alive. We compare our findings with those predicted by prevailing theories for rates of cultural change and the distribution of cultural variation.     

The sky is falling: evidence of a negativity bias in the social transmission of information

The method of serial reproduction has revealed that the social transmission of information is characterized by the gradual transformation of the original message. This transformation results from the preferential survival of certain types of information and the resolution of ambiguity. Here we present evidence of a bias favoring the social transmission of negatively-valenced information across multiple transmission episodes. Ninety-two, four-person chains transmitted a story containing unambiguously positive and unambiguously negative story events, along with ambiguous story events that could be interpreted positively or negatively. Analysis using mixed-effects modeling revealed the preferential survival of unambiguously negative events over positive events, and the increasingly negative resolution of ambiguous events across successive transmission episodes. Contrary to predictions, elevated state anxiety did not enhance the social transmission of negatively-valenced information. We also found that the survival of unambiguously negative story events was positively correlated with the negative resolution of ambiguous story events, reflecting a general negativity-bias in the social transmission of information.     

Coevolution of honest signaling and cooperative norms by cultural group selection

Evolution of cooperative norms is studied in a population where individual and group level selection are both in operation. Individuals play indirect reciprocity game within their group and follow second order norms. Individuals are norm-followers, and imitate their successful group mates. Aside from direct observation individuals can be informed about the previous actions and reputations by information transferred by others. A potential donor estimates the reputation of a potential receiver either by her own observation or by the opinion of the majority of others (indirect observation). Following a previous study (Scheuring, 2009) we assume that norms determine only the probabilities of actions, and mutants can differ in these probabilities. Similarly, we assume that individuals follow a stochastic information transfer strategy. The central question is whether cooperative norm and honest social information transfer can emerge in a population where initially only non-cooperative norms were present, and the transferred information was not sufficiently honest.It is shown that evolution can lead to a cooperative state where information transferred in a reliable manner, where generous cooperative strategies are dominant. This cooperative state emerges along a sharp transition of norms. We studied the characteristics of actions and strategies in this transition by classifying the stochastic norms, and found that a series of more and more judging strategies invade each other before the stabilization of the so-called generous judging strategy. Numerical experiments on the coevolution of social parameters (e.g. probability of direct observation and the number of indirect observers) reveal that it is advantageous to lean on indirect observation even if information transfer is much noisier than for direct observation, which is because to follow the majorities’ opinion suppresses information noise meaningfully.     

The robustness of altruism as an evolutionary strategy

Kin selection, reciprocity and group selection are widely regarded as evolutionary mechanisms capable of sustaining altruism among humans andother cooperative species. Our research indicates, however, that these mechanisms are only particular examples of a broader set of evolutionary possibilities.In this paper we present the results of a series of simple replicator simulations, run on variations of the 2–player prisoner's dilemma, designed to illustrate the wide range of scenarios under which altruism proves to be robust under evolutionary pressures. The set of mechanisms we explore is divided into four categories:correlation, group selection, imitation, and punishment. We argue that correlation is the core phenomenon at work in all four categories.     

Evolution of generous cooperative norms by cultural group selection

Evolution of cooperative norms is studied in a population where individual- and group-level selection are both in operation. Individuals play indirect reciprocity game within their group. Individuals are well informed about the previous actions and reputations, and follow second-order norms. Individuals are norm-followers, and imitate their successful group mates. In contrast to previous models where norms classify actions deterministically, we assume that norms determine only the probabilities of actions, and mutants can differ in these probabilities. The central question is how a selective cooperative norm can emerge in a population where initially only non-cooperative norms were present. It is shown that evolution leads to a cooperative state if generous cooperative strategies are dominant, although the “always defecting” and the “always cooperating”-like strategies remain stably present. The characteristics of these generous cooperative strategies and the presence of always defecting and always cooperating strategies are in concordance with experimental observations.     

A two level mutation-selection model of cultural evolution and diversity

Cultural evolution is a complex process that can happen at several levels. At the level of individuals in a population, each human bears a set of cultural traits that he or she can transmit to its offspring (vertical transmission) or to other members of his or her society (horizontal transmission). The relative frequency of a cultural trait in a population or society can thus increase or decrease with the relative reproductive success of its bearers (individual’s level) or the relative success of transmission (called the idea’s level). This article presents a mathematical model on the interplay between these two levels. The first aim of this article is to explore when cultural evolution is driven by the idea’s level, when it is driven by the individual’s level and when it is driven by both. These three possibilities are explored in relation to (a) the amount of interchange of cultural traits between individuals, (b) the selective pressure acting on individuals, (c) the rate of production of new cultural traits, (d) the individual’s capacity to remember cultural traits and to the population size. The aim is to explore the conditions in which cultural evolution does not lead to a better adaptation of individuals to the environment. This is to contrast the spread of fitness-enhancing ideas, which make individual bearers better adapted to the environment, to the spread of “selfish” ideas, which spread well simply because they are easy to remember but do not help their individual bearers (and may even hurt them). At the same time this article explores in which conditions the adaptation of individuals is maximal. The second aim is to explore how these factors affect cultural diversity, or the amount of different cultural traits in a population. This study suggests that a larger interchange of cultural traits between populations could lead to cultural evolution not improving the adaptation of individuals to their environment and to a decrease of cultural diversity.     

Cumulative cultural dynamics and the coevolution of cultural innovation and transmission: an ESS model for panmictic and structured populations

When individuals in a population can acquire traits through learning, each individual may express a certain number of distinct cultural traits. These traits may have been either invented by the individual himself or acquired from others in the population. Here, we develop a game theoretic model for the accumulation of cultural traits through individual and social learning. We explore how the rates of innovation, decay, and transmission of cultural traits affect the evolutionary stable (ES) levels of individual and social learning and the number of cultural traits expressed by an individual when cultural dynamics are at a steady‐state. We explore the evolution of these phenotypes in both panmictic and structured population settings. Our results suggest that in panmictic populations, the ES level of learning and number of traits tend to be independent of the social transmission rate of cultural traits and is mainly affected by the innovation and decay rates. By contrast, in structured populations, where interactions occur between relatives, the ES level of learning and the number of traits per individual can be increased (relative to the panmictic case) and may then markedly depend on the transmission rate of cultural traits. This suggests that kin selection may be one additional solution to Rogers's paradox of nonadaptive culture.     

An investigation of the relationship between innovation and cultural diversity

In this paper we apply reaction-diffusion models to explore the relationship between the rate of behavioural innovation and the level of cultural diversity. We investigate how both independent invention and the modification and refinement of established innovations impact on cultural dynamics and diversity. Further, we analyse these relationships in the presence of biases in cultural learning and find that the introduction of new variants typically increases cultural diversity substantially in the short term, but may decrease long-term diversity. Independent invention generally supports higher levels of cultural diversity than refinement. Repeated patterns of innovation through refinement generate characteristic oscillating trends in diversity, with increasing trends towards greater average diversity observed for medium but not low innovation rates. Conformity weakens the relationship between innovation and diversity. The level of cultural diversity, and pattern of temporal dynamics, potentially provide clues as to the underlying process, which can be used to interpret empirical data.     

Macro-evolutionary studies of cultural diversity: a review of empirical studies of cultural transmission and cultural adaptation

A growing body of theoretical and empirical research has examined cultural transmission and adaptive cultural behaviour at the individual, within-group level. However, relatively few studies have tried to examine proximate transmission or test ultimate adaptive hypotheses about behavioural or cultural diversity at a between-societies macro-level. In both the history of anthropology and in present-day work, a common approach to examining adaptive behaviour at the macro-level has been through correlating various cultural traits with features of ecology. We discuss some difficulties with simple ecological associations, and then review cultural phylogenetic studies that have attempted to go beyond correlations to understand the underlying cultural evolutionary processes. We conclude with an example of a phylogenetically controlled approach to understanding proximate transmission pathways in Austronesian cultural diversity.     

The evolution of conformist transmission in social learning when the environment changes periodically

Conformity is often observed in human social learning. Social learners preferentially imitate the majority or most common behavior in many situations, though the strength of conformity varies with the situation. Why has such a psychological tendency evolved? I investigate this problem by extending a standard model of social learning evolution with infinite environmental states (Feldman, M.W., Aoki, K., Kumm, J., 1996. Individual versus social learning: evolutionary analysis in a fluctuating environment. Anthropol. Sci. 104, 209-231) to include conformity bias. I mainly focus on the relationship between the strength of conformity bias that evolves and environmental stability, which is one of the most important factors in the evolution of social learning. Using the evolutionarily stable strategy (ESS) approach, I show that conformity always evolves when environmental stability and the cost of adopting a wrong behavior are small, though environmental stability and the cost of individual learning both negatively affect the strength of conformity.     

The minimisation of random events in the search for group structure

Summary An iterative species ranking and agglomerative clustering technique has been described. Its objectives were the minimisation of miselassification due to random species. occurrence and the demonstration of the changing role of specles in pattern formation.This work was supported by a grant from the National Science and Engineering Research Council of Canada.     

Hominid cultural transmission and the evolution of language

This paper presents the hypothesis that linguistic capacity evolved through the action of natural selection as an instrument which increased the efficiency of the cultural transmission system of early hominids. We suggest that during the early stages of hominization, hominid social learning, based on indirect social learning mechanisms and true imitation, came to constitute cumulative cultural transmission based on true imitation and the approval or disapproval of the learned behaviour of offspring. A key factor for this transformation was the development of a conceptual capacity for categorizing learned behaviour in value terms - positive or negative, good or bad. We believe that some hominids developed this capacity for categorizing behaviour, and such an ability allowed them to approve or disapprove of their offsprings- learned behaviour. With such an ability, hominids were favoured, as they could transmit to their offspring all their behavioural experience about what can and cannot be done. This capacity triggered a cultural transmission system similar to the human one, though pre-linguistic. We suggest that the adaptive advantage provided by this new system of social learning generated a selection pressure in favour of the development of a linguistic capacity allowing children to better understand the new kind of evaluative information received from parents.     

Game theory and human evolution: a critique of some recent interpretations of experimental games

Economists and psychologists have been testing Nash equilibrium predictions of game theory models of human behavior. In many instances, humans do not conform to the predictions. These results are of great interest to biologists because they also raise questions about well-known ESS models of cooperation. Cooperation in certain one-shot, anonymous interactions, and a willingness to punish others at a net cost to oneself are some of the most intriguing deviations from standard theory. One proposed explanation for these results that is receiving increasing attention invokes the cultural group selection of 'other regarding' social norms. We critically review this explanation. We conclude that experimental results reveal limits in two implicit models of cognitive structure commonly employed by economists and evolutionary biologists.