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.     

Cultural transmission can inhibit the evolution of altruistic helping

The study of culturally inherited traits has led to the suggestion that the evolution of helping behaviors is more likely with cultural transmission than without. Here we evaluate this idea through a comparative analysis of selection on helping under both genetic and cultural inheritance. We develop two simple models for the evolution of helping through cultural group selection: one in which selection on the trait depends solely on Darwinian fitness effects and one in which selection is driven by nonreproductive factors, specifically imitation of strategies achieving higher payoffs. We show that when cultural variants affect Darwinian fitness, the selection pressure on helping can be markedly increased relative to that under genetic transmission. By contrast, when variants are driven by nonreproductive factors, the selection pressure on helping may be reduced relative to that under genetic inheritance. This occurs because, unlike biological offspring, the spread of cultural variants from one group to another through imitation does not reduce the number of these variants in the source group. As a consequence, there is increased within-group competition associated with traits increasing group productivity, which reduces the benefits of helping. In these cases, selection for harming behavior (decreasing the payoff to neighbors) may occur rather than selection for helping.     

Multilevel selection and the social transmission of behavior

Many evolutionary models assume that behaviors are caused directly by genes. An implication is that behavioral uniformity should be found only in groups that are genetically uniform. Yet, the members of human social groups often behave in a uniform fashion, despite the fact that they are genetically diverse. Behavioral uniformity can occur through a variety of psychological mechanisms and social processes, such as imitation, consensus decision making, or the imposition of social norms. We present a series of models in which genes code for social transmission rules, which in turn govern the behaviors that are adopted. Transmission rules can evolve in randomly formed groups that concentrate phenotypic variation at the between-group level, favoring the evolution of altruistic behaviors and other group-advantageous traits. In addition, a direct bias toward adopting altruistic behaviors can evolve. Our models begin to show how group selection can be a strong force in human evolution, despite the absence of extreme genetic variation among groups.     

The kin composition of social groups: trading group size for degree of altruism

Why some social systems form groups composed of kin, while others do not, has gone largely untreated in the literature. Using an individual-based simulation model, we explore the demographic consequences of making kinship a criterion in group formation. We find that systems where social groups consist of one-generation breeding associations may face a serious trade-off between degree of altruism and group size that is largely mediated by their kin composition. On the one hand, restricting groups to close kin allows the evolution of highly altruistic behaviors but may limit group size to suboptimal levels, the more severely so the smaller the intrinsic fecundity of the species and the stricter the kin admission rule. Group size requirements, on the other hand, can be met by admitting nonkin into groups, but not without limiting the degree of altruism that can evolve. As a solution to this conundrum, we show that if helping roles within groups are assigned through a lottery rather than being genetically determined, maximum degrees of altruism can evolve in groups of nonrelatives of any size. Such a "lottery" mechanism may explain reproductive and helping patterns in organisms as varied as the cellular slime molds, pleometrotic ants, and Galapagos hawks.     

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.     

Why people punish defectors. Weak conformist transmission can stabilize costly enforcement of norms in cooperative dilemmas

In this paper, we present a cultural evolutionary model in which norms for cooperation and punishment are acquired via two cognitive mechanisms: (1) payoff-biased transmission-a tendency to copy the most successful individual; and (2) conformist transmission-a tendency to copy the most frequent behavior in the population. We first show that if a finite number of punishment stages is permitted (e.g. two stages of punishment occur if some individuals punish people who fail to punish non-cooperators), then an arbitrarily small amount of conformist transmission will stabilize cooperative behavior by stabilizing punishment at some n -th stage. We then explain how, once cooperation is stabilized in one group, it may spread through a multi-group population via cultural group selection. Finally, once cooperation is prevalent, we show how prosocial genes favoring cooperation and punishment may invade in the wake of cultural group selection.     

An Experimental Test of the Accumulated Copying Error Model of Cultural Mutation for Acheulean Handaxe Size

Archaeologists interested in explaining changes in artifact morphology over long time periods have found it useful to create models in which the only source of change is random and unintentional copying error, or ‘cultural mutation’. These models can be used as null hypotheses against which to detect non-random processes such as cultural selection or biased transmission. One proposed cultural mutation model is the accumulated copying error model, where individuals attempt to copy the size of another individual''s artifact exactly but make small random errors due to physiological limits on the accuracy of their perception. Here, we first derive the model within an explicit mathematical framework, generating the predictions that multiple independently-evolving artifact chains should diverge over time such that their between-chain variance increases while the mean artifact size remains constant. We then present the first experimental test of this model in which 200 participants, split into 20 transmission chains, were asked to faithfully copy the size of the previous participant''s handaxe image on an iPad. The experimental findings supported the model''s prediction that between-chain variance should increase over time and did so in a manner quantitatively in line with the model. However, when the initial size of the image that the participants resized was larger than the size of the image they were copying, subjects tended to increase the size of the image, resulting in the mean size increasing rather than staying constant. This suggests that items of material culture formed by reductive vs. additive processes may mutate differently when individuals attempt to replicate faithfully the size of previously-produced artifacts. Finally, we show that a dataset of 2601 Acheulean handaxes shows less variation than predicted given our empirically measured copying error variance, suggesting that other processes counteracted the variation in handaxe size generated by perceptual cultural mutation.     

Primate archaeology reveals cultural transmission in wild chimpanzees (Pan troglodytes verus)

Recovering evidence of past human activities enables us to recreate behaviour where direct observations are missing. Here, we apply archaeological methods to further investigate cultural transmission processes in percussive tool use among neighbouring chimpanzee communities in the Taï National Park, Côte d''Ivoire, West Africa. Differences in the selection of nut-cracking tools between neighbouring groups are maintained over time, despite frequent female transfer, which leads to persistent cultural diversity between chimpanzee groups. Through the recovery of used tools in the suggested natal territory of immigrants, we have been able to reconstruct the tool material selection of females prior to migration. In combination with direct observations of tool selection of local residents and immigrants after migration, we uncovered temporal changes in tool selection for immigrating females. After controlling for ecological differences between territories of immigrants and residents our data suggest that immigrants abandoned their previous tool preference and adopted the pattern of their new community, despite previous personal proficiency of the same foraging task. Our study adds to the growing body of knowledge on the importance of conformist tendencies in animals.     

Not-so-social learning strategies

Social learning strategies (SLSs) are rules specifying the conditions in which it would be adaptive for animals to copy the behaviour of others rather than to persist with a previously established behaviour or to acquire a new behaviour through asocial learning. In behavioural ecology, cultural evolutionary theory and economics, SLSs are studied using a ‘phenotypic gambit’—from a purely functional perspective, without reference to their underlying psychological mechanisms. However, SLSs are described in these fields as if they were implemented by complex, domain-specific, genetically inherited mechanisms of decision-making. In this article, we suggest that it is time to begin investigating the psychology of SLSs, and we initiate this process by examining recent experimental work relating to three groups of strategies: copy when alternative unsuccessful, copy when model successful and copy the majority. In each case, we argue that the reported behaviour could have been mediated by domain-general and taxonomically general psychological mechanisms; specifically, by mechanisms, identified through conditioning experiments, that make associative learning selective. We also suggest experimental manipulations that could be used in future research to resolve more fully the question whether, in non-human animals, SLSs are mediated by domain-general or domain-specific psychological mechanisms.     

Phenotypic evolution under gene-culture transmission in structured populations.

I consider a simple model for the evolution of a quantitative character is structured populations when an offspring's phenotype is determined partly by his or her genetic constitution and partly by cultural transmission of the parental phenotype. Analysis of the model indicates that when individual and group selection are in the same direction, phenotypic evolution always proceeds faster under gene-culture vs. purely genetic transmission. When individual and group selection are countervailing, altruistic characters evolve faster under gene-culture transmission when individual selection is weak and migration among groups is limited, with increased individual selection and migration tending to decrease the advantage of gene-culture transmission over purely genetic transmission. Given the prevalence of cultural transmission in higher species, these results suggest that contrary to what is often assumed, group selection may indeed by a potent evolutionary force in the evolution of altruistic characters.     

Coevolution of pathogens and cultural practices: a new look at behavioral heterogeneity in epidemics

The effect of heterogeneity within populations on the spread of infectious diseases has been a recent focus of research. Such heterogeneity may be, for example, spatial, temporal or behavioral in form. Generally, models that include population subdivision have assumed that individuals are permanently assigned to given behavioral states represented by the subpopulations. We consider a simple epidemic model in which a behavioral trait affects disease transmission, and this trait may be transferred among hosts as a consequence of social interaction. This creates a situation where the frequencies of different behavioral traits and disease states as well as their associations may change over time. We consider the impact of the culturally transmitted trait on the criterion for initial spread of the disease. We also explore the evolution of cultural traits in response to pathogen dynamics and show some conditions under which behavioral traits that reduce transmission evolve. We find that behaviors increasing the risk of infection can also evolve when they are inherently favored or when there is sufficient clustering of contacts between like behaviors.     

Review. The multiple roles of cultural transmission experiments in understanding human cultural evolution

In this paper, we explore how experimental studies of cultural transmission in adult humans can address general questions regarding the 'who, what, when and how' of human cultural transmission, and consequently inform a theory of human cultural evolution. Three methods are discussed. The transmission chain method, in which information is passed along linear chains of participants, has been used to identify content biases in cultural transmission. These concern the kind of information that is transmitted. Several such candidate content biases have now emerged from the experimental literature. The replacement method, in which participants in groups are gradually replaced or moved across groups, has been used to study phenomena such as cumulative cultural evolution, cultural group selection and cultural innovation. The closed-group method, in which participants learn in groups with no replacement, has been used to explore issues such as who people choose to learn from and when they learn culturally as opposed to individually. A number of the studies reviewed here have received relatively little attention within their own disciplines, but we suggest that these, and future experimental studies of cultural transmission that build on them, can play an important role in a broader science of cultural evolution.     

Effective population size for culturally evolving traits

Population size has long been considered an important driver of cultural diversity and complexity. Results from population genetics, however, demonstrate that in populations with complex demographic structure or mode of inheritance, it is not the census population size, N, but the effective size of a population, N_e, that determines important evolutionary parameters. Here, we examine the concept of effective population size for traits that evolve culturally, through processes of innovation and social learning. We use mathematical and computational modeling approaches to investigate how cultural N_e and levels of diversity depend on (1) the way traits are learned, (2) population connectedness, and (3) social network structure. We show that one-to-many and frequency-dependent transmission can temporally or permanently lower effective population size compared to census numbers. We caution that migration and cultural exchange can have counter-intuitive effects on N_e. Network density in random networks leaves N_e unchanged, scale-free networks tend to decrease and small-world networks tend to increase N_e compared to census numbers. For one-to-many transmission and different network structures, larger effective sizes are closely associated with higher cultural diversity. For connectedness, however, even small amounts of migration and cultural exchange result in high diversity independently of N_e. Extending previous work, our results highlight the importance of carefully defining effective population size for cultural systems and show that inferring N_e requires detailed knowledge about underlying cultural and demographic processes.     

Inclusive heritability: combining genetic and non‐genetic information to study animal behavior and culture

Phenotypic variance results from variation in biological information possessed by individuals. Quantitative geneticists often strive to partition out all environmental variance to measure heritability. Behavioral biologists and ecologists however, require methods to integrate genetic and environmental components of inherited phenotypic variance in order to estimate the evolutionary potential of traits, which encompasses any form of information that is inherited. To help develop this integration, we build on the tools of quantitative genetics and offer the concept of ‘inclusive heritability’ which identifies and unifies the various mechanisms of information transmission across generations. A controversial component of non‐genetic information is animal culture, which is the part of phenotypic variance inherited through social learning. Culture has the unique property of being transmitted horizontally and obliquely, as well as vertically. Accounting for cultural variation would allow us to examine a broader range of evolutionary mechanisms. Culture may, for instance, produce behavioral isolating mechanisms leading to speciation. To advance the study of animal culture, we offer a definition of culture that is rooted in quantitative genetics. We also offer four testable criteria to determine whether a trait is culturally inherited. These criteria may constitute a conceptual tool to study animal culture. We briefly discuss methods to partition out cultural variance. Several authors have recently called for ‘modernizing the modern synthesis’ by including non‐genetic factors such as epigenetics and phenotypic plasticity in order to more fully explain phenotypic evolution. Here, we further propose to broaden the concept of inheritance by incorporating the cultural component of behavior. Applying the concept of inclusive heritability may advance the integration of multiple forms of inheritance into the study of evolution.     

The co-evolution of individual behaviors and social institutions

We present agent-based simulations of a model of a deme-structured population in which group differences in social institutions are culturally transmitted and individual behaviors are genetically transmitted. We use a standard extended fitness accounting framework to identify the parameter space for which this co-evolutionary process generates high levels of group-beneficial behaviors. We show that intergroup conflicts may explain the evolutionary success of both: (a) altruistic forms of human sociality towards unrelated members of one's group; and (b) group-level institutional structures such as food sharing which have emerged and diffused repeatedly in a wide variety of ecologies during the course of human history. Group-beneficial behaviors may evolve if (a) they inflict sufficient fitness costs on outgroup individuals and (b) group-level institutions limit the individual fitness costs of these behaviors and thereby attenuate within-group selection against these behaviors. Thus, the evolutionary success of individually costly but group-beneficial behaviors in the relevant environments during the first 90,000 years of anatomically modern human existence may have been a consequence of distinctive human capacities in social institution building.     

Foraging behaviour alters with social environment in a juvenile songbird

Early independence from parents is a critical period where social information acquired vertically may become outdated, or conflict with new information. However, across natural populations, it is unclear if newly independent young persist in using information from parents, or if group-level effects of conformity override previous behaviours. Here, we test if wild juvenile hihi (Notiomystis cincta, a New Zealand passerine) retain a foraging behaviour from parents, or if they change in response to the behaviour of peers. We provided feeding stations to parents during chick-rearing to seed alternative access routes, and then tracked their offspring''s behaviour. Once independent, juveniles formed mixed-treatment social groups, where they did not retain preferences from their time with parents. Instead, juvenile groups converged over time to use one access route­ per group, and juveniles that moved between groups switched to copy the locally favoured option. Juvenile hihi did not copy specific individuals, even if they were more familiar with the preceding bird. Our study shows that early social experiences with parents affect initial foraging decisions, but social environments encountered later on can update transmission of arbitrary behaviours. This suggests that conformity may be widespread in animal groups, with potential cultural, ecological and evolutionary consequences.     

Selection for altruism through random drift in variable size populations

ABSTRACT: BACKGROUND: Altruistic behavior is defined as helping others at a cost to oneself and a lowered fitness. The lower fitness implies that altruists should be selected against, which is in contradiction with their widespread presence is nature. Present models of selection for altruism (kin or multilevel) show that altruistic behaviors can have 'hidden' advantages if the 'common good' produced by altruists is restricted to some related or unrelated groups. These models are mostly deterministic, or assume a frequency dependent fitness. RESULTS: Evolutionary dynamics is a competition between deterministic selection pressure and stochastic events due to random sampling from one generation to the next. We show here that an altruistic allele extending the carrying capacity of the habitat can win by increasing the random drift of "selfish" alleles. In other terms, the fixation probability of altruistic genes can be higher than those of a selfish ones, even though altruists have a smaller fitness. Moreover when populations are geographically structured, the altruists advantage can be highly amplified and the fixation probability of selfish genes can tend toward zero. The above results are obtained both by numerical and analytical calculations. Analytical results are obtained in the limit of large populations. CONCLUSIONS: The theory we present does not involve kin or multilevel selection, but is based on the existence of random drift in variable size populations. The model is a generalization of the original Fisher-Wright and Moran models where the carrying capacity depends on the number of altruists.     

Categories and gatherings: Group selection and the mythology of cultural anthropology

If the term “group selection” is to have any meaning beyond mere semantics, it must refer to situations where individuals live in groups. Although the terminology of cultural anthropology suggests that humans live in bounded and enduring gatherings that might serve as group “vehicles” of selection, we argue that none of the terms asserted to be such an entity (i.e., clans, lineages, villages, bands, tribes, populations, societies, and cultures) fulfill this requirement. This is because these terms refer to: (1) reified abstractions, (2) groups only in the sense of categories of people instead of groups in the sense of people gathered together, or (3) gatherings that are much too fluid and fuzzy in their membership to be “vehicles.” Following Murdock (1972), we refer to this obsession with groups as “anthropology's mythology,” and we suggest that it is the result of our evolved capacity for categorical perception. Although classifying phenomena into categories is useful in many situations, it has hindered our understanding of human social organization and human evolution.     

Cultural niche construction and the evolution of small family size

A model of cultural niche construction with two culturally transmitted traits is examined. The frequency of individuals with a certain general predisposition, which is transmitted vertically, plays a role as the cultural background, or the cultural niche, of the population. The cultural background determines the rate of oblique, relative to vertical, transmission of another cultural trait that affects fertility of individuals. It is assumed that individuals with fewer offspring are more likely to achieve social roles that influence the succeeding generation and are therefore overrepresented as transmitters in the process of oblique transmission. Our model suggests that even a slight overrepresentation of those with fewer offspring can drive the evolution of small family size, provided that the rate of oblique transmission depends strongly on the cultural background. In addition, our model may help to explain the time lag between the decrease in death rates and the subsequent decrease in birth rates during the demographic transition of industrializing societies.     

ALTRUISM IN MENDELIAN POPULATIONS DERIVED FROM SIBLING GROUPS: THE HAYSTACK MODEL REVISITED

A group-selection model is presented in which each group is initiated by a single fertilized female and persists for several generations before dispersal. Maynard Smith (1964) concluded that altruism could not plausibly evolve under these circumstances. I show that his conclusion is an artifact of a simplifying assumption that amounts to a worst-case scenario for group selection. When the standard donor-recipient equations for altruistic behavior are used in Maynard Smith's model, Mendelian populations derived from sibling groups are often more favorable for the evolution of altruism than are the sibling groups themselves. In general, long-term and large-scale aspects of population structure may at times be important in the evolution of altruistic and other group-advantageous behaviors.