Evolution depends on the fitness of organisms, the expected rate of reproducing. Directly getting offspring is the most basic form of fitness, but fitness can also be increased indirectly by helping genetically related individuals (such as kin) to increase their fitness. The combined effect is known as inclusive fitness. Here it is argued that a further elaboration of fitness has evolved, particularly in humans. It is called extensive fitness and it incorporates producing organisms that are merely similar in phenotype. The evolvability of this mechanism is illustrated by computations on a simple model combining heredity and behaviour. Phenotypes are driven into the direction of high fitness through a mechanism that involves an internal estimate of fitness, implicitly made within the organism itself. This mechanism has recently been conjectured to be responsible for producing agency and goals. In the model, inclusive and extensive fitness are both implemented by letting fitness increase nonlinearly with the size of subpopulations of similar heredity (for the indirect part of inclusive fitness) and of similar phenotype (for the phenotypic part of extensive fitness). Populations implementing extensive fitness outcompete populations implementing mere inclusive fitness. This occurs because groups with similar phenotype tend to be larger than groups with similar heredity, and fitness increases more when groups are larger. Extensive fitness has two components, a direct component where individuals compete in inducing others to become like them and an indirect component where individuals cooperate and help others who are already similar to them.
Cooperation is ubiquitous in every level of living organisms. It is known that spatial (network) structure is a viable mechanism for cooperation to evolve. Until recently, it has been difficult to predict whether cooperation can evolve at a network (population) level. To address this problem, Pinheiro et al. proposed a numerical metric, called Average Gradient of Selection (AGoS) in 2012. AGoS can characterize and forecast the evolutionary fate of cooperation at a population level. However, stochastic mutation of strategies was not considered in the analysis of AGoS. Here we analyzed the evolu…
We study the evolution of cooperation in a finite population interacting according to a simple model of like-with-like assortment. Evolution proceeds as a Moran process, and payoffs from the underlying cooperator-defector game are translated to positive fitnesses by an exponential transformation. These evolutionary dynamics can arise, for example, in a nest-structured population with rare migration. The use of the exponential transformation, rather than the usual linear one, is appropriate when interactions have multiplicative fitness effects, and allows for a tractable characterization of the…
It is well known that cooperation cannot be an evolutionary stable strategy for a non-iterative game in a well-mixed population. In contrast, structured populations favor cooperation since cooperators can benefit each other by forming local clusters. Previous studies have shown that scale-free networks strongly promote cooperation. However, little is known about the invasion mechanism of cooperation in scale-free networks. To study microscopic and macroscopic behaviors of cooperators' invasion, we conducted computational experiments of the evolution of cooperation in scale-free networks where,…
Inclusive Fitness Theory (IFT) was proposed half a century ago by W.D. Hamilton to explain the emergence and maintenance of cooperation between individuals that allows the existence of society. Contemporary evolutionary ecology identified several factors that increase inclusive fitness, in addition to kin-selection, such as assortation or homophily, and social synergies triggered by cooperation. Here we propose an Extend Inclusive Fitness Theory (EIFT) that includes in the fitness calculation all direct and indirect benefits an agent obtains by its own actions, and through interactions with ki…
The spatial arrangement of individuals is thought to overcome the dilemma of cooperation: When cooperators engage in clusters they might share the benefit of cooperation while being more protected against non-cooperating individuals, which benefit from cooperation but save the cost of cooperation. This is paradigmatically shown by the spatial prisoner's dilemma model. Here, we study this model in one and two spatial dimensions, but explicitly take into account that in biological setups fitness collection and selection are separated processes occurring mostly on vastly different time scales. Th…
Evolution of cooperation in the prisoner's dilemma and the public goods game is studied, where initially players belong to two independent structured populations. Simultaneously with the strategy evolution, players whose current utility exceeds a threshold are rewarded by an external link to a player belonging to the other population. Yet as soon as the utility drops below the threshold, the external link is terminated. The rewarding of current evolutionary fitness thus introduces a time-varying interdependence between the two populations. We show that, regardless of the details of the evoluti…
Cooperation is a persistent behavioral pattern of entities pooling and sharing resources. Its ubiquity in nature poses a conundrum. Whenever two entities cooperate, one must willingly relinquish something of value to the other. Why is this apparent altruism favored in evolution? Classical solutions assume a net fitness gain in a cooperative transaction which, through reciprocity or relatedness, finds its way back from recipient to donor. We seek the source of this fitness gain. Our analysis rests on the insight that evolutionary processes are typically multiplicative and noisy. Fluctuations ha…
Here, we study the evolution of specialization using realistic computer simulations of bacteria that secrete two public goods in a dynamic fluid. Through this first principles approach, we find physical factors such as diffusion, flow patterns, and decay rates are as influential as fitness economics in governing the evolution of community structure, to the extent that when mechanical factors are taken into account, (1) Generalist communities can resist becoming specialists, despite the invasion fitness of specialization, (2) Generalist and specialists can both resist cheaters despite the invas…
We compare the likelihood of different socially relevant features to allow the evolutionary emergence and maintenance of cooperation in a generalized variant of the iterated Prisoners Dilemma game. Results show that the average costs/benefit balance of cooperation is the primary constraint for its establishment and maintenance. Behavior increasing inclusive fitness such as assortation, homophily, kin-selection and tagging of individuals, is second in importance. Networks characteristics were the least important in favoring the establishment and maintenance of cooperation, despite being the mos…
In spatial evolutionary games the fitness of each individual is traditionally determined by the payoffs it obtains upon playing the game with its neighbors. Since defection yields the highest individual benefits, the outlook for cooperators is gloomy. While network reciprocity promotes collaborative efforts, chances of averting the impending social decline are slim if the temptation to defect is strong. It is therefore of interest to identify viable mechanisms that provide additional support for the evolution of cooperation. Inspired by the fact that the environment may be just as important as…
Families are places of affection and cooperation, but also of conflict. In his famous paper Parent-Offspring Conflict, Robert L. Trivers builds upon W. D. Hamilton's concept of inclusive fitness to argue for genetic conflict in parent-offspring relationships, and to derive numerical predictions on the intensity of the conflict. We propose a mathematical model of game theory that depicts how each member of a family allocates her resource budget to maximize her inclusive fitness; this latter is made of the sum of personal fitness plus the sum of relatives fitnesses weighted by Wright's coefficie…
Beijing University of Posts and Telecommunications（北京邮电大学）；Beihang University（北京航空航天大学）；Dartmouth College（达特茅斯学院）；Geisel School of Medicine at Dartmouth（达特茅斯盖泽尔医学院）；Yale University（耶鲁大学）
In human societies the probability of strategy adoption from a given person may be affected by the personal features. Now we investigate how an artificially imposed restricted ability to reproduce, overruling ones fitness, affects an evolutionary process. For this purpose we employ the evolutionary prisoner's dilemma game on different complex graphs. Reproduction restrictions can have a facilitative effect on the evolution of cooperation that sets in irrespective of particularities of the interaction network. Indeed, an appropriate fraction of less fertile individuals may lead to full supremac…
Societies consisting of cooperative individuals seem to require for their continuing success that defectors be policed. The precise connection between punishers and benefits, population structure, and division of labour, however, remains ill-understood. Many models assume costly "peer punishment" to enforce cooperation, but results in the economics literature suggest that this assumption may not be generally valid. In many human and animal societies, there is a division of labour between a purely supportive majority and a dedicated minority of police-like enforcers. Here we present several ext…
Range expansion is a universal process in biological systems, and therefore plays a part in biological evolution. Using a quantitative individual-based method based on the stochastic process, we identify that enhancing the inherent self-proliferation advantage of cooperators relative to defectors is a more effective channel to promote the evolution of cooperation in range expansion than weakening the benefit acquisition of defectors from cooperators. With this self-proliferation advantage, cooperators can rapidly colonize virgin space and establish spatial segregation more readily, which acts …
Whether or not to change strategy depends not only on the personal success of each individual, but also on the success of others. Using this as motivation, we study the evolution of cooperation in games that describe social dilemmas, where the propensity to adopt a different strategy depends both on individual fitness as well as on the strategies of neighbors. Regardless of whether the evolutionary process is governed by pairwise or group interactions, we show that plugging into the "wisdom of groups" strongly promotes cooperative behavior. The more the wider knowledge is taken into account th…
Extensive cooperation among biologically unrelated individuals is uniquely human and much current research attempts to explain this fact. We draw upon social, cultural, and psychological aspects of human uniqueness to present an integrated theory of human cooperation that explains aspects of human cooperation that are problematic for other theories (e.g., defector invasion avoidance, preferential assortment to exclude free riders, and the second order free rider problem). We propose that the evolution of human cooperative behavior required (1) a capacity for self-sustained, self-referential th…
Extensive cooperation among unrelated individuals is unique to humans, who often sacrifice personal benefits for the common good and work together to achieve what they are unable to execute alone. The evolutionary success of our species is indeed due, to a large degree, to our unparalleled other-regarding abilities. Yet, a comprehensive understanding of human cooperation remains a formidable challenge. Recent research in social science indicates that it is important to focus on the collective behavior that emerges as the result of the interactions among individuals, groups, and even societies.…
