Conditions for the evolution of altruism under darwinian selection

A model of population structure is discussed which under certain circumstances allows for evolution of altruistic traits, beyond the classical restrictions imposed by kin selection theory and related concepts such as reciprocal altruism. Essentially, the model sees a large population as socially subdivided into small groups without any barriers, however, to free random mating. An altruistic trait is defined as lowering, locally, the fitness of a carrier below that of noncarriers within the same group; but the local fitness of an individual randomly chosen in a group increases with the number of altruists. It is shown that altruism can evolve even if the groups are randomly formed. The conditions for such evolution are contrasted with those prevailing when the groups are formed either with some phenotypic assortment between the members or on the basis of kinship. It is shown that any possibility of evolution tends to rapidly disappear as groups become large, unless there is complete positive assortment or individuals in the groups are kin. The example of alarm calls is also considered, and the two extremes of random and sib-groups are contrasted, using a model by Maynard Smith. It is shown that alarm calls can evolve in small groups of unrelated individuals under conditions qualitatively similar but quantitatively more rigorous than those prevailing for sib-groups.     

The evolution of trans-generational altruism: kin selection meets niche construction

A cornerstone result of sociobiology states that limited dispersal can induce kin competition to offset the kin selected benefits of altruism. Several mechanisms have been proposed to circumvent this dilemma but all assume that actors and recipients of altruism interact during the same time period. Here, this assumption is relaxed and a model is developed where individuals express an altruistic act, which results in posthumously helping relatives living in the future. The analysis of this model suggests that kin selected benefits can then feedback on the evolution of the trait in a way that promotes altruistic helping at high rates under limited dispersal. The decoupling of kin competition and kin selected benefits results from the fact that by helping relatives living in the future, an actor is helping individuals that are not in direct competition with itself. A direct consequence is that behaviours which actors gain by reducing the common good of present and future generations can be opposed by kin selection. The present model integrates niche-constructing traits with kin selection theory and delineates demographic and ecological conditions under which altruism can be selected for; and conditions where the 'tragedy of the commons' can be reduced.     

The evolution of altruism: game theory in multilevel selection and inclusive fitness

Although the prisoner's dilemma (PD) has been used extensively to study reciprocal altruism, here we show that the n-player prisoner's dilemma (NPD) is also central to two other prominent theories of the evolution of altruism: inclusive fitness and multilevel selection. An NPD model captures the essential factors for the evolution of altruism directly in its parameters and integrates important aspects of these two theories such as Hamilton's rule, Simpson's paradox, and the Price covariance equation. The model also suggests a simple interpretation of the Price selection decomposition and an alternative decomposition that is symmetrical and complementary to it. In some situations this alternative shows the temporal changes in within- and between-group selection more clearly than the Price equation. In addition, we provide a new perspective on strong vs. weak altruism by identifying their different underlying game structures (based on absolute fitness) and showing how their evolutionary dynamics are nevertheless similar under selection (based on relative fitness). In contrast to conventional wisdom, the model shows that both strong and weak altruism can evolve in periodically formed random groups of non-conditional strategies if groups are multigenerational. An integrative approach based on the NPD helps unify different perspectives on the evolution of altruism.     

Runaway selection for cooperation and strict-and-severe punishment

Punishing defectors is an important means of stabilizing cooperation. When levels of cooperation and punishment are continuous, individuals must employ suitable social standards for defining defectors and for determining punishment levels. Here we investigate the evolution of a social reaction norm, or psychological response function, for determining the punishment level meted out by individuals in dependence on the cooperation level exhibited by their neighbors in a lattice-structured population. We find that (1) cooperation and punishment can undergo runaway selection, with evolution towards enhanced cooperation and an ever more demanding punishment reaction norm mutually reinforcing each other; (2) this mechanism works best when punishment is strict, so that ambiguities in defining defectors are small; (3) when the strictness of punishment can adapt jointly with the threshold and severity of punishment, evolution favors the strict-and-severe punishment of individuals who offer slightly less than average cooperation levels; (4) strict-and-severe punishment naturally evolves and leads to much enhanced cooperation when cooperation without punishment would be weak and neither cooperation nor punishment are too costly; and (5) such evolutionary dynamics enable the bootstrapping of cooperation and punishment, through which defectors who never punish gradually and steadily evolve into cooperators who punish those they define as defectors.     

Multilevel selection and human altruism

Views on the evolution of altruism based upon multilevel selection on structured populations pay little attention to the difference between fortuitous and deliberate processes leading to assortative grouping. Altruism may evolve when assortative grouping is fortuitously produced by forces external to the organism. But when it is deliberately produced by the same proximate mechanism that controls altruistic responses, as in humans, exploitation of altruists by selfish individuals is unlikely and altruism evolves as an individually advantageous trait. Groups formed with altruists of this sort are special, because they are not affected by subversion from within. A synergistic process where altruism is selected both at the individual and at the group level can take place.     

Adaptive evolution of social traits: origin, trajectories, and correlations of altruism and mobility

Social behavior involves "staying and helping," two individual attributes that vary considerably among organisms. Investigating the ultimate causes of such variation, this study integrates previously separate lines of research by analyzing the joint evolution of altruism and mobility. We unfold the network of selective pressures and derive how these depend on physiological costs, eco-evolutionary feedbacks, and a complex interaction between the evolving traits. Our analysis highlights habitat saturation, both around individuals (local aggregation) and around unoccupied space (local contention), as the key mediator of altruism and mobility evolution. Once altruism and mobility are allowed to evolve jointly, three general insights emerge. First, the cost of mobility affects the origin of altruism, determining whether and how quickly selfishness is overcome. Second, the cost of altruism determines which of two qualitatively different routes to sociality are taken: an evolutionary reduction of mobility, resulting in higher habitat saturation, is either preceded or followed by the adaptive rise of altruism. Third, contrary to conventional expectations, a positive correlation between evolutionarily stable levels of altruism and mobility can arise; this is expected when comparing populations that evolved under different constraints on mobility or that differ in other life-history traits.     

Levels of selection, altruism, and primate behavior.

Altruistic behaviors seem anomalous from a traditional view of Darwinian natural selection, and evolutionary explanations for them have generated much discussion. The debate centers around four major explanations: classic individual-level selection, reciprocity and game theory, kin selection, and trait-group selection. The historical context and defining criteria of each model must be reviewed before its validity can be assessed. Of these proposed mechanisms, group selection historically has been the most controversial. Although the extent to which empirical data support group selection hypotheses is uncertain, there is evidence for group-level selection among avirulent virus strains and foraging ant queens. Researchers studying mammalian behavior, particularly primatologists, have largely dismissed models of group-level selection. Most discussion of altruism among primates has focused on differences in fitness among individuals within a single group, but students of altruistic behaviors exhibited by primates also need to investigate intergroup variation with respect to these behaviors. Various altruistic behaviors are likely to have evolved through different forms of selection, and each example of apparent altruism therefore needs to be evaluated separately.     

Familial selection and the evolution of social behaviour re-examined

The equation for the fitness requirements for the evolution of altruism by diploid workers in haplodiploid species was derived from the model. of Scudo & Ghiselin (1975). When the benefit to a family is proportional to the number of altruists, the constraints on fitness were found to be the same as for haploid altruists in haplodiploid species and for workers of ether sex in diploid species, in contrast to the equation given by Scudo & Ghiselin (1975). With this correction, their results are now in agreement both with comparable allele frequency models and with kin selection games theory. The general condition for evolution of altruism under this model, when benefits are not necessarily proportional to the number of altruists, was also derived. The result appears to differ from games theory predictions, but this is solely because the basic assumptions are not comparable. Altruism is less likely to arise under these conditions, which are less favorable for altruism at gene frequencies above 0·33.     

Neocortex evolution in primates: the "social brain" is for females

According to the social intelligence hypothesis, relative neocortex size should be directly related to the degree of social complexity. This hypothesis has found support in a number of comparative studies of group size. The relationship between neocortex and sociality is thought to exist either because relative neocortex size limits group size or because a larger group size selects for a larger neocortex. However, research on primate social evolution has indicated that male and female group sizes evolve in relation to different demands. While females mostly group according to conditions set by the environment, males instead simply go where the females are. Thus, any hypothesis relating to primate social evolution has to analyse its relationship with male and female group sizes separately. Since sex-specific neocortex sizes in primates are unavailable in sufficient quantity, I here instead present results from phylogenetic comparative analyses of unsexed relative neocortex sizes and female and male group sizes. These analyses show that while relative neocortex size is positively correlated with female group size, it is negatively, or not at all correlated with male group size. This indicates that the social intelligence hypothesis only applies to female sociality.     

Inbreeding,kin selection,and primate social evolution

In this paper I argue (a) that the study of kin selection may be facilitated by looking for influences of inbreeding, which is an important aspect of a population's genetic structure; (b) that in nonhuman primates the level of inbreeding will be largely a function of the rate of migration by individuals, usually only of one sex, between social units or troops; (c) that many primate species live in relatively outbred groups, and that their social structure reflects this; and (d) that extensive social contrasts between bonnet and pigtail macaques reflect evolution by kin selection under different levels of inbreeding.     

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.     

Evolution of indirect reciprocity by social information: the role of trust and reputation in evolution of altruism

The complexity of human's cooperative behavior cannot be fully explained by theories of kin selection and group selection. If reciprocal altruism is to provide an explanation for altruistic behavior, it would have to depart from direct reciprocity, which requires dyads of individuals to interact repeatedly. For indirect reciprocity to rationalize cooperation among genetically unrelated or even culturally dissimilar individuals, information about the reputation of individuals must be assessed and propagated in a population. Here, we propose a new framework for the evolution of indirect reciprocity by social information: information selectively retrieved from and propagated through dynamically evolving networks of friends and acquaintances. We show that for indirect reciprocity to be evolutionarily stable, the differential probability of trusting and helping a reputable individual over a disreputable individual, at a point in time, must exceed the cost-to-benefit ratio of the altruistic act. In other words, the benefit received by the trustworthy must out-weigh the cost of helping the untrustworthy.     

Kin recognition and the evolution of altruism

In 1964, Hamilton formalized the idea of kin selection to explain the evolution of altruistic behaviours. Since then, numerous examples from a diverse array of taxa have shown that seemingly altruistic actions towards close relatives are a common phenomenon. Although many species use kin recognition to direct altruistic behaviours preferentially towards relatives, this important aspect of social biology is less well understood theoretically. I extend Hamilton's classic work by defining the conditions for the evolution of kin-directed altruism when recognizers are permitted to make acceptance (type I) and rejection (type II) errors in the identification of social partners with respect to kinship. The effect of errors in recognition on the evolution of kin-directed altruism depends on whether the population initially consists of unconditional altruists or non-altruists (i.e. alternative forms of non-recognizers). Factors affecting the level of these error rates themselves, their evolution and their long-term stability are discussed.     

A simple and general explanation for the evolution of altruism

We present a simple framework that highlights the most fundamental requirement for the evolution of altruism: assortment between individuals carrying the cooperative genotype and the helping behaviours of others with which these individuals interact. We partition the fitness effects on individuals into those due to self and those due to the 'interaction environment', and show that it is the latter that is most fundamental to understanding the evolution of altruism. We illustrate that while kinship or genetic similarity among those interacting may generate a favourable structure of interaction environments, it is not a fundamental requirement for the evolution of altruism, and even suicidal aid can theoretically evolve without help ever being exchanged among genetically similar individuals. Using our simple framework, we also clarify a common confusion made in the literature between alternative fitness accounting methods (which may equally apply to the same biological circumstances) and unique causal mechanisms for creating the assortment necessary for altruism to be favoured by natural selection.     

Population viscosity and the evolution of altruism

The term population viscosity refers to limited dispersal, which increases the genetic relatedness of neighbors. This effect both supports the evolution of altruism by focusing the altruists' gifts on relatives of the altruist, and also limits the extent to which altruism may emerge by exposing clusters of altruists to stiffer local competition. Previous analyses have emphasized the way in which these two effects can cancel, limiting the viability of altruism. These papers were based on models in which total population density was held fixed. We present here a class of models in which population density is permitted to fluctuate, so that patches of altruists are supported at a higher density than patches of non-altruists. Under these conditions, population viscosity can support the selection of both weak and strong altruism.     

Indirect genetic effects and inbreeding: consequences of BLUP selection for socially affected traits on rate of inbreeding

Background

Social interactions often occur among living organisms, including aquatic animals. There is empirical evidence showing that social interactions may genetically affect phenotypes of individuals and their group mates. In this context, the heritable effect of an individual on the phenotype of another individual is known as an Indirect Genetic Effect (IGE). Selection for socially affected traits may increase response to artificial selection, but also affect rate of inbreeding.

Methods

A simulation study was conducted to examine the effect of Best Linear Unbiased Prediction (BLUP) selection for socially affected traits on the rate of inbreeding. A base scenario without IGE and three alternative scenarios with different magnitudes of IGE were simulated. In each generation, 25 sires and 50 dams were mated, producing eight progeny per dam. The population was selected for 20 generations using BLUP. Individuals were randomly assigned to groups of eight members in each generation, with two families per group, each contributing four individuals. “Heritabilities” (for both direct and indirect genetic effects) were equal to 0.1, 0.3 or 0.5, and direct–indirect genetic correlations were −0.8, −0.4, 0, 0.4, or 0.8. The rate of inbreeding was calculated from generation 10 to 20.

Results

For the base scenario, the rates of inbreeding were 4.09, 2.80 and 1.95% for “heritabilities” of 0.1, 0.3 and 0.5, respectively. Overall, rates of inbreeding for the three scenarios with IGE ranged from 2.21 to 5.76% and were greater than for the base scenarios. The results show that social interaction within groups of two families increases the resemblance between estimated breeding values of relatives, which, in turn, increases the rate of inbreeding.

Conclusion

BLUP selection for socially affected traits increased the rate of inbreeding. To maintain inbreeding at an acceptable rate, a selection algorithm that restricts the increase in mean kinship, such as optimum contribution selection, is required.     

Joint evolution of multiple social traits: a kin selection analysis

General models of the evolution of cooperation, altruism and other social behaviours have focused almost entirely on single traits, whereas it is clear that social traits commonly interact. We develop a general kin-selection framework for the evolution of social behaviours in multiple dimensions. We show that whenever there are interactions among social traits new behaviours can emerge that are not predicted by one-dimensional analyses. For example, a prohibitively costly cooperative trait can ultimately be favoured owing to initial evolution in other (cheaper) social traits that in turn change the cost–benefit ratio of the original trait. To understand these behaviours, we use a two-dimensional stability criterion that can be viewed as an extension of Hamilton''s rule. Our principal example is the social dilemma posed by, first, the construction and, second, the exploitation of a shared public good. We find that, contrary to the separate one-dimensional analyses, evolutionary feedback between the two traits can cause an increase in the equilibrium level of selfish exploitation with increasing relatedness, while both social (production plus exploitation) and asocial (neither) strategies can be locally stable. Our results demonstrate the importance of emergent stability properties of multidimensional social dilemmas, as one-dimensional stability in all component dimensions can conceal multidimensional instability.     

Evidence for inbreeding depression and post-pollination selection against inbreeding in the dioecious plant Silene latifolia

In many species, inbred individuals have reduced fitness. In plants with limited pollen and seed dispersal, post-pollination selection may reduce biparental inbreeding, but knowledge on the prevalence and importance of pollen competition or post-pollination selection after non-self pollination is scarce. We tested whether post-pollination selection favours less related pollen donors and reduces inbreeding in the dioecious plant Silene latifolia. We crossed 20 plants with pollen from a sibling and an unrelated male, and with a mix of both. We found significant inbreeding depression on vegetative growth, age at first flowering and total fitness (22% in males and 14% in females). In mixed pollinations, the unrelated male sired on average 57% of the offspring. The greater the paternity share of the unrelated sire, the larger the difference in relatedness of the two males to the female. The effect of genetic similarity on paternity is consistent with predictions for post-pollination selection, although paternity, at least in some crosses, may be affected by additional factors. Our data show that in plant systems with inbreeding depression, such as S. latifolia, pollen or embryo selection after multiple-donor pollination may indeed reduce inbreeding.