Kinds of kindness: classifying the causes of altruism and cooperation.
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Altruism↗
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Results for “Evolution of cooperation”
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Cooperating for direct fitness benefits.
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Altruism↗
Cooperation in the iterated prisoner's dilemma is learned by operant conditioning mechanisms.
The prisoner's dilemma (PD) is the leading metaphor for the evolution of cooperative behavior in populations of selfish agents. Although cooperation in the iterated prisoner's dilemma (IPD) has been studied for over twenty years, most of this research has been focused on strategies that involve nonlearned behavior. Another approach is to suppose that players' selection of the preferred reply might be enforced in the same way as feeding animals track the best way to feed in changing nonstationary environments. Learning mechanisms such as operant conditioning enable animals to acquire relevant characteristics of their environment in order to get reinforcements and to avoid punishments. In this study, the role of operant conditioning in the learning of cooperation was evaluated in the PD. We found that operant mechanisms allow the learning of IPD play against other strategies. When random moves are allowed in the game, the operant learning model showed low sensitivity. On the basis of this evidence, it is suggested that operant learning might be involved in reciprocal altruism.
Animals↗
Individual variation evades the prisoner's dilemma.
BACKGROUND: The Prisoner's Dilemma (PD) is a widely used paradigm to study cooperation in evolutionary biology, as well as in fields as diverse as moral philosophy, sociology, economics and politics. Players are typically assumed to have fixed payoffs for adopting certain strategies, which depend only on the strategy played by the opponent. However, fixed payoffs are not realistic in nature. Utility functions and the associated payoffs from pursuing certain strategies vary among members of a population with numerous factors. In biology such factors include size, age, social status and expected life span; in economics they include socio-economic status, personal preference and past experience; and in politics they include ideology, political interests and public support. Thus, no outcome is identical for any two different players. RESULTS: We show that relaxing the assumption of fixed payoffs leads to frequent violations of the payoff structure required for a Prisoner's Dilemma. With variance twice the payoff interval in a linear PD matrix, for example, only 16% of matrices are valid. CONCLUSIONS: A single player lacking a valid PD matrix destroys the conditions for a Prisoner's Dilemma, so between any two players, PD games themselves are fewer still (3% in this case). This may explain why the Prisoner's Dilemma has hardly been found in nature, despite the fact that it has served as a ubiquitous (and still instructive) model in studies of the evolution of cooperation.
Analysis of Variance↗
The social evolution of bacterial pathogenesis.
Many of the genes responsible for the virulence of bacterial pathogens are carried by mobile genetic elements that can be transferred horizontally between different bacterial lineages. Horizontal transfer of virulence-factor genes has played a profound role in the evolution of bacterial pathogens, but it is poorly understood why these genes are so often mobile. Here, I present a hypothetical selective mechanism maintaining virulence-factor genes on horizontally transmissible genetic elements. For virulence factors that are secreted extracellularly, selection within hosts may favour mutant 'cheater' strains of the pathogen that do not produce the virulence factor themselves but still benefit from factors produced by other members of the pathogen population within a host. Using simple mathematical models, I show that if this occurs then selection for infectious transmission between hosts favours pathogen strains that can reintroduce functional copies of virulence-factor genes into cheaters via horizontal transfer, forcing them to produce the virulence factor. Horizontal gene transfer is thus a novel mechanism for the evolution of cooperation. I discuss predictions of this hypothesis that can be tested empirically and its implications for the evolution of pathogen virulence.
Bacteria↗
The evolution of n-player cooperation-threshold games and ESS bifurcations.
An evolutionary game of individuals cooperating to obtain a collective benefit is here modelled as an n-player Prisoner's Dilemma game. With reference to biological situations, such as group foraging, we introduce a threshold condition in the number of cooperators required to obtain the collective benefit. In the simplest version, a three-player game, complex behaviour appears as the replicator dynamics exhibits a catastrophic event separating a parameter region allowing for coexistence of cooperators and defectors and a region of pure defection. Cooperation emerges through an ESS bifurcation, and cooperators only thrive beyond a critical point in cost-benefit space. Moreover, a repelling fixed point of the dynamics acts as a barrier to the introduction of cooperation in defecting populations. The results illustrate the qualitative difference between two-player games and multiple player games and thus the limitations to the generality of conclusions from two-player games. We present a procedure to find the evolutionarily stable strategies in any n-player game with cost and benefit depending on the number of cooperators. This was previously done by Motro [1991. Co-operation and defection: playing the field and the ESS. J. Theor. Biol. 151, 145-154] in the special cases of convex and concave benefit functions and constant cost.
Animals↗
Pattern selection in a cooperative biochemical in vitro amplification system: the role of parasites.
Previously, numerical simulations have shown that evolving systems can be stabilized against emerging parasites by pattern formation in spatially extended flow reactors. Hence, it can be argued that pattern formation is a prerequisite for any experimental investigation of the biochemical evolution of cooperative function. Here, we study a model of an experimental biochemical system for the cooperative in vitro amplification of DNA strands and show that emerging parasites can induce a complex pattern formation even when no pattern formation occurs without parasites. In an adiabatic approximation where the cooperative amplification reaction is assumed to adapt fast to slowly emerging parasites, the parasite concentration itself acts as a Steuer parameter for the selection of various complex patterns. Without such an adiabatic approximation only transient patterns emerge. As any species can grow for very low concentrations, the parasite is able to infect the entire reactor and the system is finally diluted out. In the experimental biochemical system, however, the species are individual molecules and the growth of spatially separated, non-infected regions becomes feasible. Hence a cutoff threshold for the minimal concentration is applied. In these simulations the otherwise lethal infection by parasites induces the formation of spatiotemporal spirals, and this spatial structure help the host and parasitoid species to survive together. These theoretical results describe an inherent property of cooperative reactions and have an important impact on experimental investigations on the molecular evolution and complex function in spatially extended reactors. Since the formation of the complex pattern is restricted either to a rather large cutoff value or a special choice of the kinetic parameters, we, however, conclude that the persistence of evolving cooperative amplification is not possible in a simple reaction-diffusion reactor. Experimental set-ups with patchy environments, e.g. biomolecular amplification in coupled microstructured flow chambers or in microemulsion, are eligible candidates for the observation of such a self-organized pattern selection.
Animals↗
Partner's behavior, not reward distribution, determines success in an unequal cooperative task in capuchin monkeys.
It was recently demonstrated that capuchin monkeys notice and respond to distributional inequity, a trait that has been proposed to support the evolution of cooperation in the human species. However, it is unknown how capuchins react to inequitable rewards in an unrestricted cooperative paradigm in which they may freely choose both whether to participate and, within the bounds of their partner's behavior, which reward they will receive for their participation. We tested capuchin monkeys with such a design, using a cooperative barpull, which has been used with great success in the past. Contrary to our expectations, the equity of the reward distribution did not affect success or pulling behavior. However, the behavior of the partner in an unequal situation did affect overall success rates: pairs that had a tendency to alternate which individual received the higher-value food in unequal reward situations were more than twice as successful in obtaining rewards than pairs in which one individual dominated the higher-value food. This ability to equitably distribute rewards in inherently biased cooperative situations has profound implications for activities such as group hunts, in which multiple individuals work together for a single, monopolizable reward.
Animals↗
Involuntary defection and the evolutionary origins of empathy.
An occasional involuntary defection (IVD)--being unable, rather than unwilling, to donate help to others--is an intrinsic attribute of reciprocal cooperation (2003. J. Theor. Biol. 225-285). In fact, it is easy to see that--barring special circumstances--individuals that can donate help whenever requested do not need help of the same kind from others. That is, it is by no means clear why such individuals should participate in symmetric reciprocity interactions. In this paper, I analyse the consequences of introducing IVD into direct reciprocity modeling and show that a simple form of empathy--not retaliating after being punished for IVD--is a prerequisite for evolutionarily stable cooperation. Furthermore: we will see that the stability of this, empathic retaliator, strategy increases with the number of opportunities for cooperative exchanges in the life of an average individual.
Biological Evolution↗
Active linking in evolutionary games.
In the traditional approach to evolutionary game theory, the individuals of a population meet each other at random, and they have no control over the frequency or duration of interactions. Here we remove these simplifying assumptions. We introduce a new model, where individuals differ in the rate at which they seek new interactions. Once a link between two individuals has formed, the productivity of this link is evaluated. Links can be broken off at different rates. In a limiting case, the linking dynamics introduces a simple transformation of the payoff matrix. We outline conditions for evolutionary stability. As a specific example, we study the interaction between cooperators and defectors. We find a simple relationship that characterizes those linking dynamics which allow natural selection to favour cooperation over defection.
Animals↗
Punishment in animal societies.
Although positive reciprocity (reciprocal altruism) has been a focus of interest in evolutionary biology, negative reciprocity (retaliatory infliction of fitness reduction) has been largely ignored. In social animals, retaliatory aggression is common, individuals often punish other group members that infringe their interests, and punishment can cause subordinates to desist from behaviour likely to reduce the fitness of dominant animals. Punishing strategies are used to establish and maintain dominance relationships, to discourage parasites and cheats, to discipline offspring or prospective sexual partners and to maintain cooperative behaviour.
Aggression↗
Cooperation and the Prisoner's Dilemma: towards testable models of mutualism versus reciprocity
For the purpose of distinguishing between mutualism and reciprocity in nature, recent work on the evolution of cooperation has both oversimplifed and undersimplified the distinction between these two categories of cooperation. This article addresses the resulting issues of model testability, clarifies the role of time and argues that the category of 'pseudo-reciprocity' is an unnecessary complication.1997The Association for the Study of Animal Behaviour
Journal Article↗
[Evolution of mammalian sex chromosomes: cooperation of genetic and epigenetic factors].
The X and Y chromosomes of mammals, which significantly differ in structure and genetic composition, are thought to originate from a pair of autosomes. During evolution of sex chromosomes in placental mammals, the degradation of the Y chromosome and inactivation spreading along the X chromosome occurred gradually and in concert. Thus, at the molecular level, the genetic and epigenetic factors interacted toward greater differentiation of the X/Y pair. In this review, in context of a comparison permitting to trace this evolutionary pathway, we consider the structural features of mammalian sex chromosomes focusing on the X-chromosomal genes and the unique epigenetic mechanism of their regulation. Possible causes and consequences of the genes skipping inactivation and aspects of molecular mechanism of X-chromosome inactivation are discussed. A number of hypotheses are considered on evolutionary relationships of X-chromosome inactivation and other molecular processes in mammals.
Animals↗
Kin recognition: function and mechanism in avian societies.
Cooperative behaviour resulting from kin selection is widespread among animals and the ability to recognize and discriminate between kin and non-kin is a critical element in kin selection theory. Current evidence suggests that associative learning is the most likely mechanism of kin discrimination. However, surprisingly, there have been no experimental studies of the putative 'associative-learning period', the likely recognition mechanisms enabling fine discrimination between close and distant kin of similar familiarity, whether generic or individual cues are employed in kin recognition, and how recognition ability varies at different stages of a species' life history. Comparative studies of kin recognition and discrimination in cooperative and noncooperative species are also needed to shed light on the adaptive value of helping behaviour and to identify key factors in the evolution of cooperation.
Journal Article↗
Evolutionary dynamics of the continuous iterated prisoner's dilemma.
The iterated prisoner's dilemma (IPD) has been widely used in the biological and social sciences to model dyadic cooperation. While most of this work has focused on the discrete prisoner's dilemma, in which actors choose between cooperation and defection, there has been some analysis of the continuous IPD, in which actors can choose any level of cooperation from zero to one. Here, we analyse a model of the continuous IPD with a limited strategy set, and show that a generous strategy achieves the maximum possible payoff against its own type. While this strategy is stable in a neighborhood of the equilibrium point, the equilibrium point itself is always vulnerable to invasion by uncooperative strategies, and hence subject to eventual destabilization. The presence of noise or errors has no effect on this result. Instead, generosity is favored because of its role in increasing contributions to the most efficient level, rather than in counteracting the corrosiveness of noise. Computer simulation using a single-locus infinite alleles Gaussian mutation model suggest that outcomes ranging from a stable cooperative polymorphism to complete collapse of cooperation are possible depending on the magnitude of the mutational variance. Also, making the cost of helping a convex function of the amount of help provided makes it more difficult for cooperative strategies to invade a non-cooperative equilibrium, and for the cooperative equilibrium to resist destabilization by non-cooperative strategies. Finally, we demonstrate that a much greater degree of assortment is required to destabilize a non-cooperative equilibrium in the continuous IPD than in the discrete IPD. The continuous model outlined here suggests that incremental amounts of cooperation lead to rapid decay of cooperation and thus even a large degree of assortment will not be sufficient to allow cooperation to increase when cooperators are rare. The extreme degree of assortment required to destabilize the non-cooperative equilibrium, as well as the instability of the cooperative equilibrium, may help explain why cooperation in Prisoner's Dilemmas is so rare in nature.
Altruism↗
Spatialization and greater generosity in the stochastic Prisoner's Dilemma.
The iterated Prisoner's Dilemma has become the standard model for the evolution of cooperative behavior within a community of egoistic agents, frequently cited for implications in both sociology and biology. Due primarily to the work of Axelrod (1980a, 1980b, 1984, 1985), a strategy of tit for tat (TFT) has established a reputation as being particularly robust. Nowak and Sigmund (1992) have shown, however, that in a world of stochastic error or imperfect communication, it is not TFT that finally triumphs in an ecological model based on population percentages (Axelrod and Hamilton 1981), but 'generous tit for tat' (GTFT), which repays cooperation with a probability of cooperation approaching 1 but forgives defection with a probability of 1/3. In this paper, we consider a spatialized instantiation of the stochastic Prisoner's Dilemma, using two-dimensional cellular automata (Wolfram, 1984, 1986; Gutowitz, 1990) to model the spatial dynamics of populations of competing strategies. The surprising result is that in the spatial model it is not GTFT but still more generous strategies that are favored. The optimal strategy within this spatial ecology appears to be a form of 'bending over backwards', which returns cooperation for defection with a probability of 2/3--a rate twice as generous as GTFT.
Altruism↗
Synergy and discounting of cooperation in social dilemmas.
The emergence and maintenance of cooperation by natural selection is an enduring conundrum in evolutionary biology, which has been studied using a variety of game theoretical models inspired by different biological situations. The most widely studied games are the Prisoner's Dilemma, the Snowdrift game and by-product mutualism for pairwise interactions, as well as Public Goods games in larger groups of interacting individuals. Here, we present a general framework for cooperation in social dilemmas in which all the traditional scenarios can be recovered as special cases. In social dilemmas, cooperators provide a benefit to the group at some cost, while defectors exploit the group by reaping the benefits without bearing the costs of cooperation. Using the concepts of discounting and synergy for describing how benefits accumulate when more than one cooperator is present in a group of interacting individuals, we recover the four basic scenarios of evolutionary dynamics given by (i) dominating defection, (ii) coexistence of defectors and cooperators, (iii) dominating cooperation and (iv) bi-stability, in which cooperators and defectors cannot invade each other. Generically, for groups of three or more interacting individuals further, more complex, dynamics can occur. Our framework provides the first unifying approach to model cooperation in different kinds of social dilemmas.
Animals↗
Costly punishment across human societies.
Recent behavioral experiments aimed at understanding the evolutionary foundations of human cooperation have suggested that a willingness to engage in costly punishment, even in one-shot situations, may be part of human psychology and a key element in understanding our sociality. However, because most experiments have been confined to students in industrialized societies, generalizations of these insights to the species have necessarily been tentative. Here, experimental results from 15 diverse populations show that (i) all populations demonstrate some willingness to administer costly punishment as unequal behavior increases, (ii) the magnitude of this punishment varies substantially across populations, and (iii) costly punishment positively covaries with altruistic behavior across populations. These findings are consistent with models of the gene-culture coevolution of human altruism and further sharpen what any theory of human cooperation needs to explain.
Africa↗