The economics of altruism and cooperation in class-structured populations: what's in a cost? What's in a benefit?
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Altruism↗
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Results for “Evolution of cooperation”
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Image scoring and cooperation in a cleaner fish mutualism.
Humans are highly social animals and often help unrelated individuals that may never reciprocate the altruist's favour. This apparent evolutionary puzzle may be explained by the altruist's gain in social image: image-scoring bystanders, also known as eavesdroppers, notice the altruistic act and therefore are more likely to help the altruist in the future. Such complex indirect reciprocity based on altruistic acts may evolve only after simple indirect reciprocity has been established, which requires two steps. First, image scoring evolves when bystanders gain personal benefits from information gathered, for example, by finding cooperative partners. Second, altruistic behaviour in the presence of such bystanders may evolve if altruists benefit from access to the bystanders. Here, we provide experimental evidence for both of the requirements in a cleaning mutualism involving the cleaner fish Labroides dimidiatus. These cleaners may cooperate and remove ectoparasites from clients or they may cheat by feeding on client mucus. As mucus may be preferred over typical client ectoparasites, clients must make cleaners feed against their preference to obtain a cooperative service. We found that eavesdropping clients spent more time next to 'cooperative' than 'unknown cooperative level' cleaners, which shows that clients engage in image-scoring behaviour. Furthermore, trained cleaners learned to feed more cooperatively when in an 'image-scoring' than in a 'non-image-scoring' situation.
Animals↗
Anarchy, socialism and a Darwinian left.
In A Darwinian left Peter Singer aims to reconcile Darwinian theory with left wing politics, using evolutionary game theory and in particular a model proposed by Robert Axelrod, which shows that cooperation can be an evolutionarily successful strategy. In this paper I will show that whilst Axelrod's model can give support to a kind of left wing politics, it is not the kind that Singer himself envisages. In fact, it is shown that there are insurmountable problems for the idea of increasing Axelrodian cooperation within a welfare state. My surprising conclusion will be that a Darwinian left worthy of the name would be anarchistic.
Biological Evolution↗
A new niche for Vibrio logei, the predominant light organ symbiont of squids in the genus Sepiola.
Two genera of sepiolid squids--Euprymna, found primarily in shallow, coastal waters of Hawaii and the Western Pacific, and Sepiola, the deeper-, colder-water-dwelling Mediterranean and Atlantic squids--are known to recruit luminous bacteria into light organ symbioses. The light organ symbiont of Euprymna spp. is Vibrio fischeri, but until now, the light organ symbionts of Sepiola spp. have remained inadequately identified. We used a combination of molecular and physiological characteristics to reveal that the light organs of Sepiola affinis and Sepiola robusta contain a mixed population of Vibrio logei and V. fischeri, with V. logei comprising between 63 and 100% of the bacteria in the light organs that we analyzed. V. logei had not previously been known to exist in such symbioses. In addition, this is the first report of two different species of luminous bacteria co-occurring within a single light organ. The luminescence of these symbiotic V. logei strains, as well as that of other isolates of V. logei tested, is reduced when they are grown at temperatures above 20 degrees C, partly due to a limitation in the synthesis of aliphatic aldehyde, a substrate of the luminescence reaction. In contrast, the luminescence of the V. fischeri symbionts is optimal above 24 degrees C and is not enhanced by aldehyde addition. Also, V. fischeri strains were markedly more successful than V. logei at colonizing the light organs of juvenile Euprymna scolopes, especially at 26 degrees C. These findings have important implications for our understanding of the ecological dynamics and evolution of cooperative, and perhaps pathogenic, associations of Vibrio spp. with their animal hosts.
Aldehydes↗
The nature of human altruism.
Some of the most fundamental questions concerning our evolutionary origins, our social relations, and the organization of society are centred around issues of altruism and selfishness. Experimental evidence indicates that human altruism is a powerful force and is unique in the animal world. However, there is much individual heterogeneity and the interaction between altruists and selfish individuals is vital to human cooperation. Depending on the environment, a minority of altruists can force a majority of selfish individuals to cooperate or, conversely, a few egoists can induce a large number of altruists to defect. Current gene-based evolutionary theories cannot explain important patterns of human altruism, pointing towards the importance of both theories of cultural evolution as well as gene-culture co-evolution.
Altruism↗
Preferential partner selection in an evolutionary study of Prisoner's Dilemma.
Partner selection is an important process in many social interactions, permitting individuals to decrease the risks associated with cooperation. In large populations, defectors may escape punishment by roving from partner to partner, but defectors in smaller populations risk social isolation. We investigate these possibilities for an evolutionary Prisoner's Dilemma in which agents use expected payoffs to choose and refuse partners. In comparison to random or round-robin partner matching, we find that the average payoffs attained with preferential partner selection tend to be more narrowly confined to a few isolated payoff regions. Most ecologies evolve to essentially full cooperative behavior, but when agents are intolerant of defections, or when the costs of refusal and social isolation are small, we also see the emergence of wallflower ecologies in which all agents are socially isolated. Between these two extremes, we see the emergence of ecologies whose agents tend to engage in a small number of defections followed by cooperation thereafter. The latter ecologies exhibit a plethora of interesting social interaction patterns.
Algorithms↗
Epidemic spreading and cooperation dynamics on homogeneous small-world networks.
We introduce a class of small-world networks--homogeneous small-worlds--which, in contrast with the well-known Watts-Strogatz small-worlds, exhibit a homogeneous connectivity distribution, in the sense that all nodes have the same number of connections. This feature allows the investigation of pure small-world effects, detached from any associated heterogeneity. Furthermore, we use at profit the remarkable similarity between the properties of homogeneous small worlds and the heterogeneous small-worlds of Watts-Strogatz to assess the separate roles of heterogeneity and small-world effects. We investigate the dependence on these two mechanisms of the threshold for epidemic outbreaks and also of the coevolution of cooperators and defectors under natural selection. With respect to the well-studied regular homogeneous limits, we find a subtle interplay between these mechanisms. While they both contribute to reduce the threshold for an epidemic outburst, they exhibit opposite behavior in the evolution of cooperation, such that the overall results mask the true nature of their individual contribution to this process.
Journal Article↗
Cooperation among unrelated individuals: reciprocal altruism, by-product mutualism and group selection in fishes.
Cooperation among unrelated individuals can evolve not only via reciprocal altruism but also via trait-group selection or by-product mutualism (or some combination of all three categories). Therefore the (iterated) prisoner's dilemma is an insufficient paradigm for studying the evolution of cooperation. We replace this game by the cooperator's dilemma, which is more versatile because it enables all three categories of cooperative behavior to be examined within the framework of a single theory. Controlled studies of cooperation among fish provide examples of each category of cooperation. Specifically, we describe reciprocal altruism among simultaneous hermaphrodites that swap egg parcels, group-selected cooperation among fish that inspect dangerous predators and by-product mutualism in the cooperative foraging of coral-reef fish.
Altruism↗
Cooperation and competition in the evolution of ATP-producing pathways.
Heterotrophic organisms generally face a trade-off between rate and yield of adenosine triphosphate (ATP) production. This trade-off may result in an evolutionary dilemma, because cells with a higher rate but lower yield of ATP production may gain a selective advantage when competing for shared energy resources. Using an analysis of model simulations and biochemical observations, we show that ATP production with a low rate and high yield can be viewed as a form of cooperative resource use and may evolve in spatially structured environments. Furthermore, we argue that the high ATP yield of respiration may have facilitated the evolutionary transition from unicellular to undifferentiated multicellular organisms.
Adenosine Triphosphate↗
Strong reciprocity and human sociality.
Human groups maintain a high level of sociality despite a low level of relatedness among group members. This paper reviews the evidence for an empirically identifiable form of prosocial behavior in humans, which we call "strong reciprocity", that may in part explain human sociality. A strong reciprocator is predisposed to cooperate with others and punish non-cooperators, even when this behavior cannot be justified in terms of extended kinship or reciprocal altruism. We present a simple model, stylized but plausible, of the evolutionary emergence of strong reciprocity.
Altruism↗
On the dynamic persistence of cooperation: how lower individual fitness induces higher survivability.
We study a model in which cooperation and defection coexist in a dynamical steady state. In our model, subpopulations of cooperators and defectors inhabit sites on a lattice. The interactions among the individuals at a site, in the form of a prisoner's dilemma (PD) game, determine their fitnesses. The chosen PD payoff allows cooperators, but not defectors, to maintain a homogeneous population. Individuals mutate between types and migrate to neighboring sites with low probabilities. We consider both density-dependent and density-independent versions of the model. The persistence of cooperation in this model can be explained in terms of the life cycle of a population at a site. This life cycle starts when one cooperator establishes a population. Then defectors invade and eventually take over, resulting finally in the death of the population. During this life cycle, single cooperators migrate to empty neighboring sites to found new cooperator populations. The system can reach a steady state where cooperation prevails if the global "birth" rate of populations is equal to their global "death" rate. The dynamic persistence of cooperation ranges over a large section of the model's parameter space. We compare these dynamics to those from other models for the persistence of altruism and to predator-prey models.
Animals↗
Cooperation driven by mutations in multi-person Prisoner's Dilemma.
The n-person Prisoner's Dilemma is a widely used model for populations where individuals interact in groups. The evolutionary stability of populations has been analysed in the literature for the case where mutations in the population may be considered as isolated events. For this case, and assuming simple trigger strategies and many iterations per game, we analyse the rate of convergence to the evolutionarily stable populations. We find that for some values of the payoff parameters of the Prisoner's Dilemma this rate is so low that the assumption, that mutations in the population are infrequent on that time-scale, is unreasonable. Furthermore, the problem is compounded as the group size is increased. In order to address this issue, we derive a deterministic approximation of the evolutionary dynamics with explicit, stochastic mutation processes, valid when the population size is large. We then analyse how the evolutionary dynamics depends on the following factors: mutation rate, group size, the value of the payoff parameters, and the structure of the initial population. In order to carry out the simulations for groups of more than just a few individuals, we derive an efficient way of calculating the fitness values. We find that when the mutation rate per individual and generation is very low, the dynamics is characterized by populations which are evolutionarily stable. As the mutation rate is increased, other fixed points with a higher degree of cooperation become stable. For some values of the payoff parameters, the system is characterized by (apparently) stable limit cycles dominated by cooperative behaviour. The parameter regions corresponding to high degree of cooperation grow in size with the mutation rate, and in number with the group size. For some parameter values, we find more than one stable fixed point, corresponding to different structures of the initial population.
Animals↗
Coevolutionary dynamics in large, but finite populations.
Coevolving and competing species or game-theoretic strategies exhibit rich and complex dynamics for which a general theoretical framework based on finite populations is still lacking. Recently, an explicit mean-field description in the form of a Fokker-Planck equation was derived for frequency-dependent selection with two strategies in finite populations based on microscopic processes [A. Traulsen, J. C. Claussen, and C. Hauert, Phys. Rev. Lett. 95, 238701 (2005)]. Here we generalize this approach in a twofold way: First, we extend the framework to an arbitrary number of strategies and second, we allow for mutations in the evolutionary process. The deterministic limit of infinite population size of the frequency-dependent Moran process yields the adjusted replicator-mutator equation, which describes the combined effect of selection and mutation. For finite populations, we provide an extension taking random drift into account. In the limit of neutral selection, i.e., whenever the process is determined by random drift and mutations, the stationary strategy distribution is derived. This distribution forms the background for the coevolutionary process. In particular, a critical mutation rate uc is obtained separating two scenarios: above uc the population predominantly consists of a mixture of strategies whereas below uc the population tends to be in homogeneous states. For one of the fundamental problems in evolutionary biology, the evolution of cooperation under Darwinian selection, we demonstrate that the analytical framework provides excellent approximations to individual based simulations even for rather small population sizes. This approach complements simulation results and provides a deeper, systematic understanding of coevolutionary dynamics.
Animals↗
Phylogeny of cooperatively breeding cuckoos (Cuculidae, Crotophaginae) based on mitochondrial gene sequences.
The Crotophaginae is a subfamily of New World cuckoos comprising the monotypic genus Guira and three ani species ( Crotophaga). All exhibit a rare form of cooperative breeding known as plural female joint-nesting, whereby two or more females lay eggs in a single nest. I reconstructed the phylogeny of Crotophaginae using the mitochondrial genes cytochrome oxidase I, II, and III, ATPase 6 and 8, and cytochrome b. The subfamily was monophyletic, implying a single origin of cooperative breeding in New World cuckoos. Crotophaga was also monophyletic with Guira as its sister taxon. Within Crotophaga, the smooth-billed ( C. ani) and groove-billed ( C. sulcirostris) anis formed the internal clade with the greater ani ( C. major) basal to this pair. This phylogeny is consistent with differences in reproductive patterns and social organization exhibited by crotophagine cuckoos, and will serve as a framework for future study of the evolution of cooperative breeding in this subfamily.
Adenosine Triphosphatases↗
Kin discrimination and the benefit of helping in cooperatively breeding vertebrates.
In many cooperatively breeding vertebrates, a dominant breeding pair is assisted in offspring care by nonbreeding helpers. A leading explanation for this altruistic behavior is Hamilton's idea that helpers gain indirect fitness benefits by rearing relatives (kin selection). Many studies have shown that helpers typically provide care for relatives, but relatively few have shown that helpers provide closer kin with preferential care (kin discrimination), fueling the suggestion that kin selection only poorly accounts for the evolution of cooperative breeding in vertebrates. We used meta-analysis to show that (i) individuals consistently discriminate between kin, and (ii) stronger discrimination occurs in species where the benefits of helping are greater. These results suggest a general role for kin selection and that the relative importance of kin selection varies across species, as predicted by Hamilton's rule.
Altruism↗
The evolution of peptide hormones.
Despite limitations in our present knowledge it is already possible to discern the main features of peptide hormone evolution, since the same mechanisms (and indeed the same hormone molecules) function in many different ways. This underlying unity of organization has its basis in the tendency of biochemical networks, once established, to survive and diversify. The most surprising recent findings in endocrinology have been the discovery of vertebrate peptide hormones in multiple sites within the same organism, and the reports, persuasive but requiring confirmation, of vertebrate hormones in primitive unicellular organisms (20, 20a). Perhaps the major challenge for the future is to define the roles and interactions of the many peptide hormones identified in brain (18). The most primitive bacteria and the human brain, though an enormous evolutionary distance apart, may have more in common than we have recognized until now. As Axelrod & Hamilton have pointed out in a recent provocative article, "The Evolution of Cooperation" (1), bacteria, though lacking a brain, are capable of adaptive behavior that can be analysed in terms of game theory. It is clear that we can learn a great deal about the whole evolutionary process from a study of the versatile and durable peptide hormones molecules.
Animals↗
Transactional skew and assured fitness return models fail to predict patterns of cooperation in wasps.
Cooperative breeders often exhibit reproductive skew, where dominant individuals reproduce more than subordinates. Two approaches derived from Hamilton's inclusive fitness model predict when subordinate behavior is favored over living solitarily. The assured fitness return (AFR) model predicts that subordinates help when they are highly likely to gain immediate indirect fitness. Transactional skew models predict dominants and subordinates "agree" on a level of reproductive skew that induces subordinates to join groups. We show the AFR model to be a special case of transactional skew models that assumes no direct reproduction by subordinates. We use data from 11 populations of four wasp species (Polistes, Liostenogaster) as a test of whether transactional frameworks suffice to predict when subordinate behavior should be observed in general and the specific level of skew observed in cooperative groups. The general prediction is supported; in 10 of 11 cases, transactional models correctly predict presence or absence of cooperation. In contrast, the specific prediction is not consistent with the data. Where cooperation occurs, the model accurately predicts highly biased reproductive skew between full sisters. However, the model also predicts that distantly related or unrelated females should cooperate with low skew. This prediction fails: cooperation with high skew is the observed norm. Neither the generalized transactional model nor the special-case AFR model can explain this significant feature of wasp sociobiology. Alternative, nontransactional hypotheses such as parental manipulation and kin recognition errors are discussed.
Animals↗
Cost minimization by helpers in cooperative vertebrates.
When parents invest heavily in reproduction they commonly suffer significant energetic costs. Parents reduce the long-term fitness implications of these costs through increased foraging and reduced reproductive investment in the future. Similar behavioral modifications might be expected among helpers in societies of cooperative vertebrates, in which helping is associated with energetic costs. By using multivariate analyses and experiments, we show that in cooperative meerkats, Suricata suricatta, helping is associated with substantial short-term growth costs but limited long-term fitness costs. This association forms because individual contributions to cooperation are initially condition dependent, and, because when helpers invest heavily in cooperation, they increase their foraging rate during the subsequent nonbreeding period and reduce their level of cooperative investment in the subsequent reproductive period. These results provide a unique demonstration that despite significant short-term costs, helpers, like breeders, are able to reduce the fitness consequences of these costs through behavioral modifications.
Animals↗