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The evolution of bourgeois, parasitic, and cooperative reproductive behaviors in fishes.

Among vertebrate classes, fishes exhibit by far the greatest variability in competitive and cooperative behaviors in male reproduction. Scramble competition between reproductive males is one possibility. Another possibility occurs when resources, mates, or locations can be monopolized, in which case males may invest in primary access to fertilizations by adopting a "bourgeois" strategy, or they may employ alternative mating tactics to evade the reproductive monopoly of other males. Adaptations in morphology, physiology, and behavior to bourgeois and alternative phenotypes are highly divergent. Here I review the functional characteristics that differ between bourgeois and parasitic phenotypes, and discuss the variability of alternative reproductive tactics at the levels of plasticity, determination, and selection. Examples will illustrate the importance of ecology, and will suggest that variation in reproductive tactics is largely adaptive. Behavioral solutions to competition for mates and fertilizations often involve agonistic behavior and conflict, but also cooperation among competitors (e.g., when subordinate males pay a price to bourgeois males for gaining access to fertilizable eggs). Application of molecular genetic tools has helped to uncover intricate sexual and social relationships in various fish species, including species that display some of the most complex reproductive and social patterns known among the vertebrates.

Agonistic Behavior↗

PubMed2005

Altruistic punishment and the origin of cooperation.

How did human cooperation evolve? Recent evidence shows that many people are willing to engage in altruistic punishment, voluntarily paying a cost to punish noncooperators. Although this behavior helps to explain how cooperation can persist, it creates an important puzzle. If altruistic punishment provides benefits to nonpunishers and is costly to punishers, then how could it evolve? Drawing on recent insights from voluntary public goods games, I present a simple evolutionary model in which altruistic punishers can enter and will always come to dominate a population of contributors, defectors, and nonparticipants. The model suggests that the cycle of strategies in voluntary public goods games does not persist in the presence of punishment strategies. It also suggests that punishment can only enforce payoff-improving strategies, contrary to a widely cited "folk theorem" result that suggests that punishment can allow the evolution of any strategy.

Altruism↗

PubMed2003

Indirect reciprocity among imperfect individuals.

Reciprocal cooperation occurs when the overall benefits of receiving help exceed the costs of donating help (Q. Rev. Biol. 46 (197) 35). That is, individuals in good condition--for whom the pertinent costs are relatively small; donate help in order to secure reciprocity in their hour of need--when the benefits of receiving a donation are large. Consequently, reciprocity occurs among individuals who occasionally need help. In particular, such individuals will be unable to help others, no matter how deserving, when in need of help themselves--involuntary defection. This paper deals with the effects of involuntary defection in the context of a specific model of indirect reciprocity (i.e. reciprocal altruism that is directed toward all the cooperative members of the community) due to Nowak and Sigmund (J. Theor. Biol.194 (1998b) 561: Sections 2-4). In that model, the authors formulate the decision rules for conditional cooperation in the context of indirect reciprocity, and demonstrate that these decision rules can account for a long-term persistence of cooperation. Here we show that addition of involuntary defection to the decision rules formulated by Nowak and Sigmund results in indirect reciprocity that is evolutionary stable under appropriate conditions. Moreover, for a wide range of parameter values, evolutionary stability of cooperation requires a mixture of conditional- and unconditional-altruist behaviors. To recollect, unconditional altruist strategy can be viewed as conditional altruist strategy sans the ability to decide when the help-soliciting individual should be refused help. That is, given involuntary defection, stability of cooperation requires an occasional forgiveness, if only by default, of a failure to donate help. Thus, we see that evolutionary stable indirect reciprocity does not require perfection in either the ability to assess the merits of the help-soliciting individuals, or the ability to donate help when it is merited. On the contrary, we are forced to conclude that reciprocity, at least in the current case, is stable only among imperfect individuals.

Altruism↗

PubMed2006

A generalized discrete model linking rippling pattern formation and individual cell reversal statistics in colonies of myxobacteria.

Self-organization processes in multicellular aggregates of bacteria and amoebae offer fascinating insights into the evolution of cooperation and differentiation of cells. During myxobacterial development a variety of spatio-temporal patterns emerges such as counterpropagating waves of cell density that are known as rippling. Recently, several models have been introduced that qualitatively reproduce these patterns. All models include active motion and a collision-triggered reversal of individual bacteria. Here, we present a systematic study of a generalized discrete model that is based on similar assumptions as the continuous model by Igoshin et al (2001 Proc. Natl Acad. Sci. USA 98 14913). We find counterpropagating as well as unidirectional rippling waves in extended regions of the parameter space. If the interaction strength and the degree of cooperativity are large enough, rippling patterns appear even in the absence of a refractory period. We show for the first time that the experimentally observed double peak in the reversal statistics of bacteria in rippling colonies (Welch and Kaiser 2001 Proc. Natl Acad. Sci. USA 98 14907) can be reproduced in simulations of counterpropagating rippling waves which are dominant in experiments. In addition, the reversal statistics in the pre-rippling phase is correctly reproduced.

Biological Evolution↗

PubMed1994

Kin selection, social structure, gene flow, and the evolution of chimpanzees.

Hypotheses about chimpanzee social behavior, phylogeography, and evolution were evaluated by noninvasive genotyping of free-ranging individuals from 20 African sites. Degrees of relatedness among individuals in one community were inferred from allele-sharing at eight nuclear simple sequence repeat (SSR) loci. Males are related on the order of half-siblings, and homozygosity is significantly increased at several SSR loci compared to Hardy-Weinberg expectations. These data support the kin-selection hypothesis for the evolution of cooperation among males. Sequence variation patterns at two mitochondrial loci indicate historically high long-distance gene flow and clarify the relationships among three allopatric subspecies. The unexpectedly large genetic distance between the western subspecies, Pan troglodytes verus, and the other two subspecies suggests a divergence time of about 1.58 million years. This result, if confirmed at nuclear loci and supported by eco-behavioral data, implies that P. t. verus should be elevated to full species rank.

Africa↗

PubMed1996

Evolutionary stability in the n-person iterated prisoner's dilemma.

The iterated prisoner's dilemma game has been used extensively in the study of the evolution of cooperative behaviours in social and biological systems. The concept of evolutionary stability provides a useful tool to analyse strategies for playing the game. Most results on evolutionary stability, however, are based on the 2-person iterated prisoner's dilemma game. This paper extends the results in the 2-person game and shows that no finite mixture of pure strategies in the n-person iterated prisoner's dilemma game can be evolutionarily stable, where n > 2. The paper also shows that evolutionary stability can be achieved if mistakes are allowed in the n-person game.

Animals↗

PubMed2006

Relationship between dilemma occurrence and the existence of a weakly dominant strategy in a two-player symmetric game.

Defining the dilemma game by the proposition, A game cannot sustain an increase of cooperation strategy in its strategy distribution, we deduced that the substance of a dilemma can be expressed by a productive summation of the static factor and the dynamic factor independently. A static factor is an element of the game's structure that influences a possible dilemma, which relates to a game's structural deviation from a situation where the cooperation strategy can be weakly dominant over other strategies. In contrast, a dynamic factor refers to a strategy distribution's influence on the dilemma by affecting the game dynamics. In a 2x2 game, the existence of a dilemma can be determined only by a static factor. That is, whether or not a dilemma occurs is related only to the structural effect of the game. On the other hand, in a more-than-two-strategies game, both static and dynamic factors determine the occurrence of a dilemma, and the static factor cannot solely explain the occurrence of a dilemma.

Algorithms↗

PubMed2006

The group covariance effect and fitness trade-offs during evolutionary transitions in individuality.

Transforming our understanding of life is the realization that evolution occurs not only among individuals within populations but also through the integration of groups of preexisting individuals into a new higher-level individual, that is, through evolutionary transitions in individuality. During evolutionary transitions (such as during the origin of gene networks, bacteria-like cells, eukaryotic cells, multicellular organisms, and societies), fitness must be reorganized; specifically, it must be transferred from the lower- to the higher-level units and partitioned among the lower-level units that specialize in the fitness components of the new higher-level individual. This paper studies the role of fitness trade-offs in fitness reorganization, the evolution of cooperation, and the conversion of a group into a new individual during the origin of multicellular life. Specifically, this study shows that the fitness of the group is augmented over the average fitness of its members according to a covariance effect. This covariance effect appears to be one of the first emergent properties of the group and a general aspect of groups with multiplicative properties that are themselves averages of properties of lower-level units. The covariance effect allows groups to break through the constraints that govern their members, and this effect likely applies to group dynamics in other fields.

Animals↗

PubMed2005

Altruism may arise from individual selection.

The fact that humans cooperate with non-kin in large groups, or with people they will never meet again, is a long-standing evolutionary puzzle. Altruism, the capacity to perform costly acts that confer benefits on others, is at the core of cooperative behavior. Behavioral experiments show that humans have a predisposition to cooperate with others and to punish non-cooperators at personal cost (so-called strong reciprocity) which, according to standard evolutionary game theory arguments, cannot arise from selection acting on individuals. This has led to the suggestion of group and cultural selection as the only mechanisms that can explain the evolutionary origin of human altruism. We introduce an agent-based model inspired on the Ultimatum Game, that allows us to go beyond the limitations of standard evolutionary game theory and show that individual selection can indeed give rise to strong reciprocity. Our results are consistent with the existence of neural correlates of fairness and in good agreement with observations on humans and monkeys.

Altruism↗

PubMed2004

Variation in behaviour promotes cooperation in the Prisoner's Dilemma game.

The Prisoner's Dilemma game is widely used to investigate how cooperation between unrelated individuals can evolve by natural selection. In this game, each player can either 'cooperate' (invest in a common good) or 'defect' (exploit the other's investment). If the opponent cooperates, you get R if you cooperate and T if you defect. If the opponent defects, you get S if you cooperate and P if you defect. Here T > R > 0 and P > S, so that 'defect' is the best response to any action by the opponent. Thus in a single play of the game, each player should defect. In our game, a fixed maximum number of rounds of the Prisoner's Dilemma game is played against the same opponent. A standard argument based on working backwards from the last round shows that defection on all rounds is the only stable outcome. In contrast, we show that if extrinsic factors maintain variation in behaviour, high levels of co-operation are stable. Our results highlight the importance of extrinsic variability in determining the outcome of evolutionary games.

Biological Evolution↗

PubMed2001

Fundamental clusters in spatial 2x2 games.

The notion of fundamental clusters is introduced, serving as a rule of thumb to characterize the statistical properties of the complex behaviour of cellular automata such as spatial 2 x 2 games. They represent the smallest cluster size determining the fate of the entire system. Checking simple growth criteria allows us to decide whether the cluster-individuals, e.g. some mutant family, are capable of surviving and invading a resident population. In biology, spatial 2 x 2 games have a broad spectrum of applications ranging from the evolution of cooperation and intraspecies competition to disease spread. This methodological study allows simple classifications and long-term predictions in various biological and social models to be made. For minimal neighbourhood types, we show that the intuitive candidate, a 3 x 3 cluster, turns out to be fundamental with certain weak limitations for the Moore neighbourhood but not for the Von Neumann neighbourhood. However, in the latter case, 2 x 2 clusters generally serve as reliable indicators to whether a strategy survives. Stochasticity is added to investigate the effects of varying fractions of one strategy present at initialization time and to discuss the rich dynamic properties in greater detail. Finally, we derive Liapunov exponents for the system and show that chaos reigns in a small region where the two strategies coexist in dynamical equilibrium.

Animals↗

PubMed2006

The costs and benefits of genetic heterogeneity in resistance against parasites in social insects.

The occurrence of polygyny and polyandry in social insects has long puzzled evolutionary biologists. If cooperation requires genetic relatedness, how do we explain the occurrence and maintenance of mechanisms that reduce the degree of relatedness among colony members? A much-discussed hypothesis states that genetically diverse colonies are more resistant to parasitism than homogenous colonies because genetic diversity reduces the spread of a disease within a colony. However, as we will argue in this note, a necessary condition for the parasite hypothesis is that genetically heterogeneous colonies have a larger suite of parasites that are capable of infecting them. This implicit relationship is important because it implies that even if the cost per infection is reduced, this may not be sufficient to offset the increased rate of acquiring infections. The advantages of genetic heterogeneity as a defense against parasites thus may not be as big as commonly thought.

Animals↗

PubMed1993

The role of autoshaping in cooperative two-player games between starlings.

We report a study of the behavior of starlings in laboratory situations inspired by the "prisoner's dilemma." Our purpose is to investigate some possible mechanisms for the maintenance of cooperation by reciprocity and to investigate the process of autoshaping at a trial-by-trial level. In Experiment 1, pairs of starlings housed in adjacent cages played a discrete-trial "game" in which food could be obtained only by "cooperation." In this game, pecking at a response key eliminated the opportunity to obtain food but produced food for the partner. If neither bird pecked, neither had the opportunity to obtain food in that trial. Some level of cooperation persisted for several sessions whether the birds had been pretrained for a high or low probability of pecking at the key. The probability of a cooperative response was higher after trials in which the partner responded (and a reward was obtained) than after trials in which neither bird responded (and no reward was obtained), but the probability of a response was even higher after trials in which the same bird had responded, even though no reward was obtained by the actor in these trials. This behavior did not require visual presence of another player, because similar results were obtained in Experiment 2 (a replicate of Experiment 1 in which the members of the pair could not see each other) and in Experiment 3, a game in which each starling played with a computer responding with "tit for tat." Using an omission schedule, in which food was given in all trials in which the bird did not peck, Experiment 4 showed that pecking could be maintained by autoshaping. In this experiment, overall probability of pecking decreased with experience, due to a drop in the tendency to peck in consecutive trials. The probability of pecking in trials following a reinforced trial did not decrease with experience. An implementation of the Rescorla-Wagner model for this situation was capable of reproducing molar, but not molecular, aspects of our results. The results violate the predictions of several game-theoretical models for the evolution of cooperation, including tit for tat, generous tit for tat, and the superior win-stay-lose-shift.

Journal Article↗

PubMed2002

Conflicting levels of selection in the accumulation of mitochondrial defects in Saccharomyces cerevisiae.

The somatic accumulation of defective mitochondria causes human degenerative syndromes, senescence in fungi, and male sterility in plants. These diverse phenomena may result from conflicts between natural selection at different levels of organization. Such conflicts are fundamental to the evolution of cooperating groups, from cells to populations. We present a model in which defective mitochondrial genomes accumulate because of a within-cell replication advantage when among-cell selection for efficient respiration is relaxed. We tested the model by using experimental populations of the yeast Saccharomyces cerevisiae. We constructed yeast strains that were heteroplasmic for mitochondrial mutations that destroy the ability to respire (the petite phenotype) and followed the accumulation of mitochondrial defects in cultures with different effective population sizes. As predicted by the model, the inability to respire evolved only in small populations of S. cerevisiae, where among-cell selection favoring cells that can respire was reduced relative to within-cell selection favoring parasitic mitochondria. In a control experiment, mitochondrial point mutations that confer resistance to chloramphenicol showed no tendency to change in frequency under any culture conditions. The accumulation of some mitochondrial defects is therefore an evolutionary process, involving multiple levels of selection. The relative intensities of within- and among-cell selection may also explain the tissue specificity of human mitochondrial defects.

Cell Respiration↗

PubMed2003

Give unto others: genetically unrelated cotton-top tamarin monkeys preferentially give food to those who altruistically give food back.

Altruistic food giving among genetically unrelated individuals is rare in nature. The few examples that exist suggest that when animals give food to unrelated others, they may do so on the basis of mutualistic or reciprocally altruistic relationships. We present the results of four experiments designed to tease apart the factors mediating food giving among genetically unrelated cotton-top tamarins (Saguinus oedipus), a cooperatively breeding New World primate. In experiment 1 we show that individuals give significantly more food to a trained conspecific who unilaterally gives food than to a conspecific who unilaterally never gives food. The apparent contingency of the tamarins' food-giving behaviour motivated the design of experiments 2-4. Results from all three experiments show that altruistic food giving is mediated by prior acts of altruistic food giving by a conspecific. Specifically, tamarins do not give food to unrelated others when the food received in the past represents the by-product of another's selfish actions (experiments 2 and 3) or when a human experimenter gives them food (experiment 4) as did the unilateral altruist in experiment 1. By contrast, if one tamarin gives another food without obtaining any immediate benefit, then the recipient is more likely to give food in return. Overall, results show that tamarins altruistically give food to genetically unrelated conspecifics, discriminate between altruistic and selfish actions, and give more food to those who give food back. Tamarins therefore have the psychological capacity for reciprocally mediated altruism.

Animals↗

PubMed1993

Friendship and the evolution of co-operation.

Recent theoretical research on the repeated Prisoner's Dilemma game (PD) suggests that the evolution of co-operative behaviour depends on the rate at which social groups break apart and new groups form. Long-lasting social groups tend to favour the evolution of co-operation. It is therefore of interest to construct models in which individuals must choose whether to maintain their current social relationships. This is done in the present paper, and the analysis suggests that unco-operative individuals will seek to move quickly from one social relationship to another, while co-operative individuals will seek to maintain relationships with other co-operative individuals. As a result of their differing decisions about changing social relationships, co-operators tend to interact with other co-operators to a greater extent than do unco-operative individuals, and this difference is an important determinant of the results of the model. In contrast to the original analyses of the repeated Prisoner's Dilemma, the model presented here allows for the existence of stable equilibria with both co-operative and unco-operative individuals simultaneously present in the same population. It is proved that the position of these equilibria (and hence the frequency of co-operators) depends on the size of the various payoffs that define the Prisoner's Dilemma game. In addition, it is proved that co-operation can be maintained at a high frequency so long as co-operators are able to maintain their relationships with other co-operators over long periods of time. This is true even if unco-operative individuals are able to move from one relationship to another at a very high rate.

Biological Evolution↗

PubMed2002

Evolving the division of labour: generalists, specialists and task allocation.

The evolutionary dynamics of specialization, in the context of the division of labour, are investigated. Individuals associate in groups in which benefits are shared and costs borne individually; each individual is either a generalist who can perform all the necessary tasks, a specialist who performs a sub-set of the necessary tasks, or a parasite who contributes nothing to the group. The implications of the model are explored analytically and through both numerical and Monte Carlo methods. These methods demonstrate the evolution of populations towards stable arrangements of specialists and generalists. The fittest populations are those that divide tasks fairly and associate in large, highly specialized groups. Generalists have a distinct advantage in small groups, but the presence of generalists, ironically, lowers group fitness. Parasites are able to invade both specialized and non-specialized populations. A basic model for the continuous division of labour is also presented, demonstrating a tendency for populations to evolve increasingly unfair divisions of labour. This last result implies that an evolutionary ratchet favours disparity between the workload of specialist populations.

Animals↗

PubMed2001

Evolution of altruism in stepping-stone populations with overlapping generations.

We study the evolution of altruism in one- and two-dimensional stepping-stone populations with discrete overlapping generations. We find that increasing survival probability facilitates the evolution of altruism, in agreement with recent results for a patch-structured population. We allow the altruistic behaviour to affect either fecundity or survival probability. In the first case, altruism is favoured compared to a randomly interacting population, but in the second case, altruism is less likely to evolve. We consider the iterated prisoner's dilemma as a description of an altruistic interaction and compare our results with recent simulations of lattice populations.

Altruism↗