Cooperative Phenotype Research Articles

The importance of need-altruism and kin-altruism to blood donor behaviour for black and white people.

Need-altruism (a preference to help people in need) and kin-altruism (a preference to help kin over non-kin) underlie two hypotheses for voluntary blood donation: (i) Need-altruism underlies motivations for volunteer blood donation and (ii) Black people express a stronger preference for kin-altruism, which is a potential barrier to donation. This paper tests these hypotheses and explores how need- and kin-altruism are associated with wider altruistic motivations, barriers, and strategies to encourage donation. We assessed need- and kin-altruism, other mechanisms-of-altruism (e.g., reluctant-altruism), barriers, strategies to encourage donation, donor status, and willingness-to-donate across four groups based on ethnicity (Black; White), nationality (British; Nigerian), and country-of-residence: (i) Black-British people (n = 395), and Black-Nigerian people (ii) in the UK (n = 97) or (iii) across the rest of the world (n = 101), and (v) White-British people in the UK (n = 452). We also sampled a Black-Nigerian Expert group (n = 60). Need-altruism was higher in donors and associated with willingness-to-donate in non-donors. Levels of kin-altruism did not differ between Black and White people, but need-altruism was lower in Black-British people. Kin-altruism was associated with a preference for incentives, and need-altruism with a preference for recognition (e.g., a thank you) as well as an increased willingness-to-donate for Black non-donors. Need-altruism underlies a blood-donor-cooperative-phenotype. Need-altruism is central to blood donation, in particular recruitment. Lower need-altruism may be a specific barrier for Black-British people. Kin-altruism is important for Black non-donors. The blood donor cooperative phenotype deserves further consideration. Implications for blood services are discussed.

A model for the dissemination of circulating tumour cell clusters involving platelet recruitment and a plastic switch between cooperative and individual behaviours

BackgroundIn spite of extensive research, cancer remains a major health problem worldwide. As cancer progresses, cells acquire traits that allow them to disperse and disseminate to distant locations in the body – a process known as metastasis. While in the vasculature, these cells are referred to as circulating tumour cells (CTCs) and can manifest either as single cells or clusters of cells (i.e., CTC clusters), with the latter being the most aggressive. The increased metastatic potential of CTC clusters is generally associated with cooperative group benefits in terms of survival, including increased resistance to shear stress, anoikis, immune attacks and drugs. However, the adoption of a group phenotype poses a challenge when exiting the vasculature (extravasation) as the large size can hinder the passage through vessel walls. Despite their significant role in the metastatic process, the mechanisms through which CTC clusters extravasate remain largely unknown. Based on the observed in vivo association between CTC clusters and platelets, we hypothesized that cancer cells take advantage of the platelet-derived Transforming Growth Factor Beta 1 (TGF-β1) – a signalling factor that has been widely implicated in many aspects of cancer, to facilitate their own dissemination. To address this possibility, we evaluated the effect of exogenous TGF-β1 on an experimentally evolved non-small cell lung cancer cell line that we previously developed and used to investigate the biology of CTC clusters.ResultsWe found that exogenous TGF-β1 induced the dissociation of clusters in suspension into adherent single cells. Once adhered, cells released their own TGF-β1 and were able to individually migrate and invade in the absence of exogenous TGF-β1. Based on these findings we developed a model that involves a TGF-β1-mediated plastic switch between a cooperative phenotype and a single-celled stage that enables the extravasation of CTC clusters.ConclusionsThis model allows for the possibility that therapies can be developed against TGF-β1 signalling components and/or TGF-β1 target genes to suppress the metastatic potential of CTC clusters. Considering the negative impact that metastasis has on cancer prognosis and the lack of therapies against this process, interfering with the ability of CTC clusters to switch between cooperative and individual behaviours could provide new strategies to improve patient survival.

Cooperative phenotype predicts climate change belief and pro-environmental behaviour

Understanding the psychological causes of variation in climate change belief and pro-environmental behaviour remains an urgent challenge for the social sciences. The “cooperative phenotype” is a stable psychological preference for cooperating in social dilemmas that involve a tension between individual and collective interest. Since climate change poses a social dilemma on a global scale, this issue may evoke similar psychological processes as smaller social dilemmas. Here, we investigate the relationships between the cooperative phenotype and climate change belief and behaviour with a representative sample of New Zealanders (N = 897). By linking behaviour in a suite of economic games to self-reported climate attitudes, we show robust positive associations between the cooperative phenotype and both climate change belief and pro-environmental behaviour. Furthermore, our structural equation models support a motivated reasoning account in which the relationship between the cooperative phenotype and pro-environmental behaviour is mediated by climate change belief. These findings suggest that common psychological mechanisms underlie cooperation in both micro-scale social dilemmas and larger-scale social dilemmas like climate change.

Advertising cooperative phenotype through costly signals facilitates collective action.

Around the world, people engage in practices that involve self-inflicted pain and apparently wasted resources. Researchers theorized that these practices help stabilize within-group cooperation by assorting individuals committed to collective action. While this proposition was previously studied using existing religious practices, we provide a controlled framework for an experimental investigation of various predictions derived from this theory. We recruited 372 university students in the Czech Republic who were randomly assigned into either a high-cost or low-cost condition and then chose to play a public goods game (PGG) either in a group that wastes money to signal commitment to high contributions in the game or to play in the group without such signals. We predicted that cooperators would assort in the high-cost revealed group and that, despite these costs, they would contribute more to the common pool and earn larger individual rewards over five iterations of PGG compared with the concealed group and participants in the low-cost condition. The results showed that the assortment of cooperators was more effective in the high-cost condition and translated into larger contributions of the remaining endowment to the common pool, but participants in the low-cost revealed group earned the most. We conclude that costly signals can serve as an imperfect assorting mechanism, but the size of the costs needs to be carefully balanced with potential benefits to be profitable.

Collapse of Intra-Tumor Cooperation Induced by Engineered Defector Cells.

Simple SummaryUsing experiments in vitro and a mathematical model, I show that genetically modified cancer cells in which the gene for an essential growth factor is knocked out can spread in a population of cells that produce that growth factor, leading to an overall reduction in proliferation—a proof of principle for a potential treatment that harnesses clonal selection by impairing intra-tumor cooperation.Anti-cancer therapies promote clonal selection of resistant cells that evade treatment. Effective therapy must be stable against the evolution of resistance. A potential strategy based on concepts from evolutionary game theory is to impair intra-tumor cooperation using genetically modified cells in which genes coding for essential growth factors have been knocked out. Such engineered cells would spread by clonal selection, driving the collapse of intra-tumor cooperation and a consequent reduction in tumor growth. Here, I test this idea in vitro in four cancer types (neuroendocrine pancreatic cancer, mesothelioma, lung adenocarcinoma and multiple myeloma). A reduction, or even complete eradication, of the producer clone and the consequent reduction in cell proliferation, is achieved in some but not all cases by introducing a small fraction of non-producer cells in the population. I show that the collapse of intra-tumor cooperation depends on the cost/benefit ratio of growth factor production. When stable cooperation among producer and non-producer cells occurs, its collapse can be induced by increasing the number of growth factors available to the cells. Considerations on nonlinear dynamics in the framework of evolutionary game theory explain this as the result of perturbation of the equilibrium of a system that resembles a public goods game, in which the production of growth factors is a cooperative phenotype. Inducing collapse of intra-tumor cooperation by engineering cancer cells will require the identification of growth factors that are essential for the tumor and that have a high cost of production for the cell.

Evolution of non-kin cooperation: social assortment by cooperative phenotype in guppies.

Cooperation among non-kin constitutes a conundrum for evolutionary biology. Theory suggests that non-kin cooperation can evolve if individuals differ consistently in their cooperative phenotypes and assort socially by these, such that cooperative individuals interact predominantly with one another. However, our knowledge of the role of cooperative phenotypes in the social structuring of real-world animal populations is minimal. In this study, we investigated cooperative phenotypes and their link to social structure in wild Trinidadian guppies (Poecilia reticulata). We first investigated whether wild guppies are repeatable in their individual levels of cooperativeness (i.e. have cooperative phenotypes) and found evidence for this in seven out of eight populations, a result which was mostly driven by females. We then examined the social network structure of one of these populations where the expected fitness impact of cooperative contexts is relatively high, and found assortment by cooperativeness, but not by genetic relatedness. By contrast, and in accordance with our expectations, we did not find assortment by cooperativeness in a population where the expected fitness impact of cooperative contexts is lower. Our results provide empirical support for current theory and suggest that assortment by cooperativeness is important for the evolution and persistence of non-kin cooperation in real-world populations.

Extending the Cooperative Phenotype: Assessing the Stability of Cooperation across Countries.

This paper studies whether individual cooperation is stable across settings and over time. Involving more than 7,000 subjects on two different continents, this study documents positive correlation in cooperative behavior across economic games in Norway, Sweden, Austria, and the United States. The game measures also correlate with a tendency to make deontological judgments in moral dilemmas, and display of general trust toward strangers. Using time-variation in the data, we test whether temporal stability of behavior is similar in the United States and Norway, and find similar stability estimates for both the American and Norwegian samples. The findings here provide further evidence of the existence of a stable behavioral inclination toward prosociality – a “cooperative phenotype,” as it has recently been termed. Also in line with previous research, we find that punishment and cooperation seem to be uncorrelated.

By genes alone: a model selectionist argument for genetical explanations of cooperation in non-human organisms

I distinguish two versions of kin selection theory—a purely genetic version (GKST) and a version that also appeals to cultural (i.e. non-genetically-derived) forms of cooperation (WKST)—and present an argument in favor of using the former when it comes to accounting for the evolution of cooperation in non-human organisms. Specifically, I first show that both GKST and WKST are equally mathematically coherent—they can both be derived from the Price equation—but not necessarily equally empirically plausible, as they are based on different assumptions about the inheritance system underlying the cooperative phenotype. Given this, I then, second, present a model selection theoretic argument in favor of GKST over WKST. This argument is based on the fact that, in non-human cases, the former theory is likely to be as empirically successful as WKST, while containing fewer degrees of freedom. I end by defending both the intrinsic importance of this argument and its relevance to the discussion surrounding the “gene’s eye view of evolution.”

Environmental modification via a quorum sensing molecule influences the social landscape of siderophore production.

Bacteria produce a wide variety of exoproducts that favourably modify their environment and increase their fitness. These are often termed ‘public goods’ because they are costly for individuals to produce and can be exploited by non-producers (cheats). The outcome of conflict over public goods is dependent upon the prevailing environment and the phenotype of the individuals in competition. Many bacterial species use quorum sensing (QS) signalling molecules to regulate the production of public goods. QS, therefore, determines the cooperative phenotype of individuals, and influences conflict over public goods. In addition to their regulatory functions, many QS molecules have additional properties that directly modify the prevailing environment. This leads to the possibility that QS molecules could influence conflict over public goods indirectly through non-signalling effects, and the impact of this on social competition has not previously been explored. The Pseudomonas aeruginosa QS signal molecule PQS is a powerful chelator of iron which can cause an iron starvation response. Here, we show that PQS stimulates a concentration-dependent increase in the cooperative production of iron scavenging siderophores, resulting in an increase in the relative fitness of non-producing siderophore cheats. This is likely due to an increased cost of siderophore output by producing cells and a concurrent increase in the shared benefits, which accrue to both producers and cheats. Although PQS can be a beneficial signalling molecule for P. aeruginosa, our data suggest that it can also render a siderophore-producing population vulnerable to competition from cheating strains. More generally, our results indicate that the production of one social trait can indirectly affect the costs and benefits of another social trait.

The foundress’s dilemma: group selection for cooperation among queens of the harvester ant, Pogonomyrmex californicus

The evolution of cooperation is a fundamental problem in biology, especially for non-relatives, where indirect fitness benefits cannot counter within-group inequalities. Multilevel selection models show how cooperation can evolve if it generates a group-level advantage, even when cooperators are disadvantaged within their group. This allows the possibility of group selection, but few examples have been described in nature. Here we show that group selection can explain the evolution of cooperative nest founding in the harvester ant Pogonomyrmex californicus. Through most of this species’ range, colonies are founded by single queens, but in some populations nests are instead founded by cooperative groups of unrelated queens. In mixed groups of cooperative and single-founding queens, we found that aggressive individuals had a survival advantage within their nest, but foundress groups with such non-cooperators died out more often than those with only cooperative members. An agent-based model shows that the between-group advantage of the cooperative phenotype drives it to fixation, despite its within-group disadvantage, but only when population density is high enough to make between-group competition intense. Field data show higher nest density in a population where cooperative founding is common, consistent with greater density driving the evolution of cooperative foundation through group selection.

The Good, the Bad, and the Unflinchingly Selfish: Cooperative Decision-Making Can Be Predicted with High Accuracy Using Only Three Behavioral Types

The human willingness to pay costs to benefit anonymous others is often explained by social preferences: rather than only valuing their own material payoff, people also care in some fashion about the outcomes of others. But how successful is this concept of outcome-based social preferences for actually predicting out-of-sample behavior? We investigate this question by having 1067 human subjects each make 20 cooperation decisions, and using machine learning to predict their last 5 choices based on their first 15. We find that decisions can be predicted with high accuracy by models that include outcome-based features and allow for heterogeneity across individuals in baseline cooperativeness and the weights placed on the outcome-based features (AUC=0.89). It is not necessary, however, to have a fully heterogeneous model -- excellent predictive power (AUC=0.88) is achieved by a model that allows three different sets of baseline cooperativeness and feature weights (i.e. three behavioral types), defined based on the participant's cooperation frequency in the 15 training trials: those who cooperated at least half the time, those who cooperated less than half the time, and those who never cooperated. Finally, we provide evidence that this inclination to cooperate cannot be well proxied by other personality/morality survey measures or demographics, and thus is a natural kind (or cooperative phenotype)

Humans display a 'cooperative phenotype' that is domain general and temporally stable.

Understanding human cooperation is of major interest across the natural and social sciences. But it is unclear to what extent cooperation is actually a general concept. Most research on cooperation has implicitly assumed that a person's behaviour in one cooperative context is related to their behaviour in other settings, and at later times. However, there is little empirical evidence in support of this assumption. Here, we provide such evidence by collecting thousands of game decisions from over 1,400 individuals. A person's decisions in different cooperation games are correlated, as are those decisions and both self-report and real-effort measures of cooperation in non-game contexts. Equally strong correlations exist between cooperative decisions made an average of 124 days apart. Importantly, we find that cooperation is not correlated with norm-enforcing punishment or non-competitiveness. We conclude that there is a domain-general and temporally stable inclination towards paying costs to benefit others, which we dub the 'cooperative phenotype'.

The Cooperative Phenotype

Understanding human cooperation is of major interest across the natural and social sciences. But it is unclear to what extent cooperation is actually a general concept. Most research on cooperation has implicitly assumed that a person’s behavior in one cooperative context is related to their behavior in other settings, and at later times. However, there is little empirical evidence in support of this assumption. Here, we provide such evidence by collecting thousands of game decisions from over 1,400 individuals. A person’s decisions in different cooperation games are correlated, as are those decisions and both self-report and real-effort measures of cooperation in non-game contexts. Equally strong correlations exist between cooperative decisions made an average of 124 days apart. Importantly, we find that cooperation is not correlated with norm-enforcing punishment or non-competitiveness. We conclude that there is a domain-general and temporally stable inclination toward paying costs to benefit others, which we dub the ‘cooperative phenotype’.

A fitness trade-off between local competition and dispersal in Vibrio cholerae biofilms

Bacteria commonly grow in densely populated surface-bound communities, termed biofilms, where they gain benefits including superior access to nutrients and resistance to environmental insults. The secretion of extracellular polymeric substances (EPS), which bind bacterial collectives together, is ubiquitously associated with biofilm formation. It is generally assumed that EPS secretion is a cooperative phenotype that benefits all neighboring cells, but in fact little is known about the competitive and evolutionary dynamics of EPS production. By studying Vibrio cholerae biofilms in microfluidic devices, we show that EPS-producing cells selectively benefit their clonemates and gain a dramatic advantage in competition against an isogenic EPS-deficient strain. However, this advantage carries an ecological cost beyond the energetic requirement for EPS production: EPS-producing cells are impaired for dispersal to new locations. Our study establishes that a fundamental tradeoff between local competition and dispersal exists among bacteria. Furthermore, this tradeoff can be governed by a single phenotype.

Cooperative Adaptation to Establishment of a Synthetic Bacterial Mutualism

To understand how two organisms that have not previously been in contact can establish mutualism, it is first necessary to examine temporal changes in their phenotypes during the establishment of mutualism. Instead of tracing back the history of known, well-established, natural mutualisms, we experimentally simulated the development of mutualism using two genetically-engineered auxotrophic strains of Escherichia coli, which mimic two organisms that have never met before but later establish mutualism. In the development of this synthetic mutualism, one strain, approximately 10 hours after meeting the partner strain, started oversupplying a metabolite essential for the partner's growth, eventually leading to the successive growth of both strains. This cooperative phenotype adaptively appeared only after encountering the partner strain but before the growth of the strain itself. By transcriptome analysis, we found that the cooperative phenotype of the strain was not accompanied by the local activation of the biosynthesis and transport of the oversupplied metabolite but rather by the global activation of anabolic metabolism. This study demonstrates that an organism has the potential to adapt its phenotype after the first encounter with another organism to establish mutualism before its extinction. As diverse organisms inevitably encounter each other in nature, this potential would play an important role in the establishment of a nascent mutualism in nature.

Social network reciprocity as a phase transition in evolutionary cooperation

In evolutionary dynamics the understanding of cooperative phenomena in natural and social systems has been the subject of intense research during decades. We focus attention here on the so-called "lattice reciprocity" mechanisms that enhance evolutionary survival of the cooperative phenotype in the prisoner's dilemma game when the population of Darwinian replicators interact through a fixed network of social contacts. Exact results on a "dipole model" are presented, along with a mean-field analysis as well as results from extensive numerical Monte Carlo simulations. The theoretical framework used is that of standard statistical mechanics of macroscopic systems, but with no energy considerations. We illustrate the power of this perspective on social modeling, by consistently interpreting the onset of lattice reciprocity as a thermodynamical phase transition that, moreover, cannot be captured by a purely mean-field approach.

Collective action in an RNA virus

Abstract A recent empirical study by Turner and Chao on the evolution of competitive interactions among phage virus strains revealed that a strain grown at high rates of co-infection evolved towards lowered fitness relative to an ancestral strain. The authors went on to show that the fitness pay-off matrix between the evolved and ancestral strain conforms to the prisoners’ dilemma. In this paper, I use Turner and Chao’s data to parameterize a simple model of parasite collective action. The prisoners’ dilemma is based on pairwise interactions of a discrete cooperate/defect nature. In contrast, the collective action model explicitly deals with individual–group interactions where the extent of cooperation is a continuous variable. I argue here that the ‘collective action’ modelling approach is more appropriate than the prisoners’ dilemma for the biology of virus evolution, and hence better able to form a predictive framework for further work on related strains of virus, linking mixing ecology, cooperative phenotype and fitness. Furthermore, the collective action model is used to motivate discussion on the evolutionary ecology of viruses, with a focus on the ‘levels of selection’ debate and the evolution of virulence.