Confusion cannot explain cooperative behavior in public goods games.

Some scholars find that behavioral variation in the public goods game is explained by variations in participants' understanding of how to maximize payoff and that confusion leads to cooperation. Their findings lead them to question the common assumption in behavioral economics experiments that choices reflect motivations. We conduct two experiments, in which we minimize confusion by providing participants with increased training. We also introduce a question that specifically assesses participants' understanding of payoff maximization choices. Our experimental results show that the distribution of behavior types is significantly different when participants play with computers versus humans. A significant increase in contributions is also observed when participants play with humans compared to when they play with computers. Moreover, social norms may be the main motive for contributions when playing with computers. Our findings suggest that social preferences, rather than confusion, play a crucial role in determining contributions in public goods games when playing with humans. We therefore argue that the assumption in behavioral economics experiments that choices reveal motivations is indeed valid.

Parallel ecological and evolutionary responses to selection in a natural bacterial community.

Evolution can occur over ecological timescales, suggesting a potentially important role for rapid evolution in shaping community trait distributions. However, evidence of concordant eco-evolutionary dynamics often comes from in vitro studies of highly simplified communities, and measures of ecological and evolutionary dynamics are rarely directly comparable. Here, we quantified how ecological species sorting and rapid evolution simultaneously shape community trait distributions by tracking within- and between-species changes in a key trait in a complex bacterial community. We focused on the production of siderophores; bacteria use these costly secreted metabolites to scavenge poorly soluble iron and to detoxify environments polluted with toxic nonferrous metals. We found that responses to copper-imposed selection within and between species were ultimately the same-intermediate siderophore levels were favored-and occurred over similar timescales. Despite being a social trait, this level of siderophore production was selected regardless of whether species evolved in isolation or in a community context. Our study suggests that evolutionary selection can play a pivotal role in shaping community trait distributions within natural, highly complex, bacterial communities. Furthermore, trait evolution may not always be qualitatively affected by interactions with other community members.

Human Prosocial Preferences Are Related to Slow-Wave Activity in Sleep.

Prosocial behavior is crucial for the smooth functioning of the society. Yet, individuals differ vastly in the propensity to behave prosocially. Here, we try to explain these individual differences under normal sleep conditions without any experimental modulation of sleep. Using a portable high-density EEG, we measured the sleep data in 54 healthy adults (28 females) during a normal night's sleep at the participants' homes. To capture prosocial preferences, participants played an incentivized public goods game in which they faced real monetary consequences. The whole-brain analyses showed that a higher relative slow-wave activity (SWA, an indicator of sleep depth) in a cluster of electrodes over the right temporoparietal junction (TPJ) was associated with increased prosocial preferences. Source localization and current source density analyses further support these findings. Recent sleep deprivation studies imply that sleeping enough makes us more prosocial; the present findings suggest that it is not only sleep duration, but particularly sufficient sleep depth in the TPJ that is positively related to prosociality. Because the TPJ plays a central role in social cognitive functions, we speculate that sleep depth in the TPJ, as reflected by relative SWA, might serve as a dispositional indicator of social cognition ability, which is reflected in prosocial preferences. These findings contribute to the emerging framework explaining the link between sleep and prosocial behavior by shedding light on the underlying mechanisms.

Social evolution of shared biofilm matrix components.

Biofilm formation is an important and ubiquitous mode of growth among bacteria. Central to the evolutionary advantage of biofilm formation is cell-cell and cell-surface adhesion achieved by a variety of factors, some of which are diffusible compounds that may operate as classical public goods-factors that are costly to produce but may benefit other cells. An outstanding question is how diffusible matrix production, in general, can be stable over evolutionary timescales. In this work, using Vibrio cholerae as a model, we show that shared diffusible biofilm matrix proteins are indeed susceptible to cheater exploitation and that the evolutionary stability of producing these matrix components fundamentally depends on biofilm spatial structure, intrinsic sharing mechanisms of these components, and flow conditions in the environment. We further show that exploitation of diffusible adhesion proteins is localized within a well-defined spatial range around cell clusters that produce them. Based on this exploitation range and the spatial distribution of cell clusters, we constructed a model of costly diffusible matrix production and related these length scales to the relatedness coefficient in social evolution theory. Our results show that production of diffusible biofilm matrix components is evolutionarily stable under conditions consistent with natural biofilm habitats and host environments. We expect the mechanisms revealed in this study to be relevant to other secreted factors that operate as cooperative public goods in bacterial communities and the concept of exploitation range and the associated analysis tools to be generally applicable.

The restart effect in social dilemmas shows humans are self-interested not altruistic.

Do economic games show evidence of altruistic or self-interested motivations in humans? A huge body of empirical work has found contrasting results. While many participants routinely make costly decisions that benefit strangers, consistent with the hypothesis that humans exhibit a biologically novel form of altruism (or "prosociality"), many participants also typically learn to pay fewer costs with experience, consistent with self-interested individuals adapting to an unfamiliar environment. Key to resolving this debate is explaining the famous "restart effect," a puzzling enigma whereby failing cooperation in public goods games can be briefly rescued by a surprise restart. Here we replicate this canonical result, often taken as evidence of uniquely human altruism, and show that it 1) disappears when cooperation is invisible, meaning individuals can no longer affect the behavior of their groupmates, consistent with strategically motivated, self-interested, cooperation; and 2) still occurs even when individuals are knowingly grouped with computer players programmed to replicate human decisions, consistent with confusion. These results show that the restart effect can be explained by a mixture of self-interest and irrational beliefs about the game's payoffs, and not altruism. Consequently, our results suggest that public goods games have often been measuring self-interested but confused behaviors and reject the idea that conventional theories of evolution cannot explain the results of economic games.

Kin selection for cooperation in natural bacterial populations.

Bacteria produce a range of molecules that are secreted from the cell and can provide a benefit to the local population of cells. Laboratory experiments have suggested that these "public goods" molecules represent a form of cooperation, favored because they benefit closely related cells (kin selection). However, there is a relative lack of data demonstrating kin selection for cooperation in natural populations of bacteria. We used molecular population genetics to test for signatures of kin selection at the genomic level in natural populations of the opportunistic pathogen Pseudomonas aeruginosa We found consistent evidence from multiple traits that genes controlling putatively cooperative traits have higher polymorphism and greater divergence and are more likely to harbor deleterious mutations relative to genes controlling putatively private traits, which are expressed at similar rates. These patterns suggest that cooperative traits are controlled by kin selection, and we estimate that the relatedness for social interactions in P. aeruginosa is r = 0.84. More generally, our results demonstrate how molecular population genetics can be used to study the evolution of cooperation in natural populations.

Cue-driven microbial cooperation and communication: evolving quorum sensing with honest signaling.

BACKGROUND: Quorum sensing (QS) is the ability of microorganisms to assess local clonal density by measuring the extracellular concentration of signal molecules that they produce and excrete. QS is also the only known way of bacterial communication that supports the coordination of within-clone cooperative actions requiring a certain threshold density of cooperating cells. Cooperation aided by QS communication is sensitive to cheating in two different ways: laggards may benefit from not investing in cooperation but enjoying the benefit provided by their cooperating neighbors, whereas Liars explicitly promise cooperation but fail to do so, thereby convincing potential cooperating neighbors to help them, for almost free. Given this double vulnerability to cheats, it is not trivial why QS-supported cooperation is so widespread among prokaryotes. RESULTS: We investigated the evolutionary dynamics of QS in populations of cooperators for whom the QS signal is an inevitable side effect of producing the public good itself (cue-based QS). Using spatially explicit agent-based lattice simulations of QS-aided threshold cooperation (whereby cooperation is effective only above a critical cumulative level of contributions) and three different (analytical and numerical) approximations of the lattice model, we explored the dynamics of QS-aided threshold cooperation under a feasible range of parameter values. We demonstrate three major advantages of cue-driven cooperation. First, laggards cannot wipe out cooperation under a wide range of reasonable environmental conditions, in spite of an unconstrained possibility to mutate to cheating; in fact, cooperators may even exclude laggards at high cooperation thresholds. Second, lying almost never pays off, if the signal is an inevitable byproduct (i.e., the cue) of cooperation; even very cheap fake signals are selected against. And thirdly, QS is most useful if local cooperator densities are the least predictable, i.e., if their lattice-wise mean is close to the cooperation threshold with a substantial variance. CONCLUSIONS: Comparing the results of the four different modeling approaches indicates that cue-driven threshold cooperation may be a viable evolutionary strategy for microbes that cannot keep track of past behavior of their potential cooperating partners, in spatially viscous and in well-mixed environments alike. Our model can be seen as a version of the famous greenbeard effect, where greenbeards coexist with defectors in a evolutionarily stable polymorphism. Such polymorphism is maintained by the condition-dependent trade-offs of signal production which are characteristic of cue-based QS.

Ecological dependencies and the illusion of cooperation in microbial communities.

Ecological dependencies - where organisms rely on other organisms for survival - are a ubiquitous feature of life on earth. Multicellular hosts rely on symbionts to provide essential vitamins and amino acids. Legume plants similarly rely on nitrogen-fixing rhizobia to convert atmospheric nitrogen to ammonia. In some cases, dependencies can arise via loss-of-function mutations that allow one partner to benefit from the actions of another. It is common in microbiology to label ecological dependencies between species as cooperation - making it necessary to invoke cooperation-specific frameworks to explain the phenomenon. However, in many cases, such traits are not (at least initially) cooperative, because they are not selected for because of the benefits they confer on a partner species. In contrast, dependencies in microbial communities may originate from fitness benefits gained from genomic-streamlining (i.e. Black Queen Dynamics). Here, we outline how the Black Queen Hypothesis predicts the formation of metabolic dependencies via loss-of-function mutations in microbial communities, without needing to invoke any cooperation-specific explanations. Furthermore we outline how the Black Queen Hypothesis can act as a blueprint for true cooperation as well as discuss key outstanding questions in the field. The nature of interactions in microbial communities can predict the ability of natural communities to withstand and recover from disturbances. Hence, it is vital to gain a deeper understanding of the factors driving these dynamic interactions over evolutionary time.

Plasmid-free cheater cells commonly evolve during laboratory growth.

It has been nearly a century since the isolation and use of penicillin, heralding the discovery of a wide range of different antibiotics. In addition to clinical applications, such antibiotics have been essential laboratory tools, allowing for selection and maintenance of laboratory plasmids that encode cognate resistance genes. However, antibiotic resistance mechanisms can additionally function as public goods. For example, extracellular beta-lactamases produced by resistant cells that subsequently degrade penicillin and related antibiotics allow neighboring plasmid-free susceptible bacteria to survive antibiotic treatment. How such cooperative mechanisms impact selection of plasmids during experiments in laboratory conditions is poorly understood. Here, we show in multiple bacterial species that the use of plasmid-encoded beta-lactamases leads to significant curing of plasmids in surface-grown bacteria. Furthermore, such curing was also evident for aminoglycoside phosphotransferase and tetracycline antiporter resistance mechanisms. Alternatively, antibiotic selection in liquid growth led to more robust plasmid maintenance, although plasmid loss was still observed. The net outcome of such plasmid loss is the generation of a heterogenous population of plasmid-containing and plasmid-free cells, leading to experimental confounds that are not widely appreciated.IMPORTANCEPlasmids are routinely used in microbiology as readouts of cell biology or tools to manipulate cell function. Central to these studies is the assumption that all cells in an experiment contain the plasmid. Plasmid maintenance in a host cell typically depends on a plasmid-encoded antibiotic resistance marker, which provides a selective advantage when the plasmid-containing cell is grown in the presence of antibiotic. Here, we find that growth of plasmid-containing bacteria on a surface and to a lesser extent in liquid culture in the presence of three distinct antibiotic families leads to the evolution of a significant number of plasmid-free cells, which rely on the resistance mechanisms of the plasmid-containing cells. This process generates a heterogenous population of plasmid-free and plasmid-containing bacteria, an outcome which could confound further experimentation.

Selective pressures for public antibiotic resistance.

The rapid increase of antibiotic-resistant pathogens is severely limiting our current treatment possibilities. An important subset of the resistance mechanisms conferring antibiotic resistance have public effects, allowing otherwise susceptible bacteria to also survive antibiotic treatment. As susceptible bacteria can survive treatment without bearing the metabolic cost of producing the resistance mechanism, there is potential to increase their relative frequency in the population and, as such, select against resistant bacteria. Multiple studies showed that this altered selection for resistance is dependent on various environmental and treatment parameters. In this review, we provide a comprehensive overview of their most important findings and describe the main factors impacting the selection for resistance. In-depth understanding of the driving forces behind selection can aid in the design and implementation of alternative treatments which limit the risk of resistance development.

Cooperation in public goods game does not require assortment and depends on population density.

The threshold public goods game is one of the best-known models of non-linear public goods dilemmas. Cooperators and defectors typically coexist in this game when the population is assumed to follow the so-called structured deme model. In this article, we develop a dynamical model of a general N-player game in which there is no deme structure: Individuals interact with randomly chosen neighbours and selection occurs between randomly chosen pairs of individuals. We show that in the deterministic limit, the dynamics in this model leads to the same replicator dynamics as in the structured deme model, i.e., coexistence of cooperators and defectors is typical in threshold public goods game even when the population is completely well mixed. We extend the model to study the effect of density dependence and density fluctuation on the dynamics. We show analytically and numerically that decreasing population density increases the equilibrium frequency of cooperators till the fixation of this strategy, but below a critical density cooperators abruptly disappear from the population. Our numerical investigations show that weak density fluctuations enhance cooperation, while strong fluctuations suppress it.

Antisocial peer exclusion does not eliminate the effectiveness of prosocial peer exclusion in structured populations.

While prosocial exclusion has been proposed as a mechanism to maintain cooperation in one-shot social dilemma games, the evolution of prosocial peer exclusion in response to the threat of antisocial peer exclusion, particularly in structured populations, remains insufficiently understood. In this study, we employ an extended spatial public goods game to investigate the evolution of prosocial peer exclusion and its impact on cooperation in the presence of both prosocial and antisocial peer exclusion. Our model encompasses four primary strategies: traditional cooperation and defection, prosocial peer exclusion targeting defectors, and antisocial peer exclusion targeting cooperators. Our findings illuminate that the presence of antisocial peer exclusion significantly disrupts network reciprocity and suppresses cooperation. However, when coexisting with prosocial peer exclusion, it does not undermine the latter's efficacy in upholding cooperation, except in scenarios with low exclusion costs Unlike the cooperation-sustaining cyclic dominance pattern observed in the exclusive presence of prosocial peer exclusion, the co-presence of prosocial and antisocial peer exclusion gives rise to more intricate pathways for maintaining cooperation. These pathways include cyclic dominance involving traditional cooperation, prosocial peer exclusion, and antisocial peer exclusion, or a similar pattern involving traditional defection and the two exclusion strategies, or even cyclic dominance among all four strategies. In essence, our study enhances the theoretical framework concerning the effectiveness of the prosocial exclusion strategy, contributing to a more comprehensive understanding of its dynamics.

Bidding against the wind: A choice experiment in green energy, green jobs and offshore views in North Carolina, USA.

Offshore wind development is in its nascent stages in the United States. Recent research indicates that the visual impacts of offshore wind farms are viewed negatively by the general population. This North Carolina application is the first US-focused discrete choice experiment that explicitly asks respondents to consider the positive local and global benefits from offshore wind development, such as job creation and greenhouse gas emission reductions, simultaneously with their visual impacts. We find significant willingness to pay (WTP) for reducing the visual impacts of offshore wind farms, and that the extent of disamenity varies in the population and with placement along developed tourist towns (as much as $783/year for three years) or preserved coastlines (as much as $451/year for three years). We also find that some preference classes value projects that create permanent jobs and reduce carbon emissions. We use our estimates of preferences for the positive and negative attributes to explore specific wind farm configurations and locations that could achieve positive consensus in a heterogenous population.

Factors Influencing the Perceived Economic Benefits of Innovative Agri-Environmental Contracts.

Continued innovation in contract design may enhance the delivery of agri-environmental climate public goods (AECPG), but barriers to adoption arise in terms of how farmers perceive the economic benefits. Therefore, this paper examines survey data from Ireland and the Netherlands to analyse whether land managers agree that results-based, collective action, value chain and land tenure contracts for the delivery of AECPG are understandable, applicable to their farm and economically beneficial. Using Probit models, we also identify groups of land managers who perceive the different contract types as being economically beneficial, and these findings can inform policymakers of farmer groups who need adequate consideration during the design of agri-environmental contracts. For example, greater incentives could encourage older farmers to enrol in results-based contracts in Ireland and value chain contracts in the Netherlands. We also find a link between contract duration and the perceived economic benefits of collective action contracts in both countries, with land managers in Ireland desiring a longer duration. We highlight that policymakers and land managers in Ireland could apply lessons from the design of agri-environmental contracts in the Netherlands, where they are more common and varied. Greater knowledge exchange between users and non-users of such contracts would also help bridge the gap between theory and practice in both countries.

Multiple roles of released c-type cytochromes in tuning electron transport and physiological status of Geobacter sulfurreducens.

Dissimilatory metal-reducing bacteria (DMRB) can transfer electrons to extracellular insoluble electron acceptors and play important roles in geochemical cycling, biocorrosion, environmental remediation, and bioenergy generation. c-type cytochromes (c-Cyts) are synthesized by DMRB and usually transported to the cell surface to form modularized electron transport conduits through protein assembly, while some of them are released as extracellularly free-moving electron carriers in growth to promote electron transport. However, the type of these released c-Cyts, the timing of their release, and the functions they perform have not been unrevealed yet. In this work, after characterizing the types of c-Cyts released by Geobacter sulfurreducens under a variety of cultivation conditions, we found that these c-Cyts accumulated up to micromolar concentrations in the surrounding medium and conserved their chemical activities. Further studies demonstrated that the presence of c-Cyts accelerated the process of microbial extracellular electron transfer and mediated long-distance electron transfer. In particular, the presence of c-Cyts promoted the microbial respiration and affected the physiological state of the microbial community. In addition, c-Cyts were observed to be adsorbed on the surface of insoluble electron acceptors and modify electron acceptors. These results reveal the overlooked multiple roles of the released c-Cyts in acting as public goods, delivering electrons, modifying electron acceptors, and even regulating bacterial community structure in natural and artificial environments.

The insurance role in workplace health promotion: a comparative analysis of the United States and the Philippines.

There is a void in the health promotion literature concerning the role of health insurance. This study offers a comparative analysis of employee health insurance in the United States and the Philippines. As the first study of its kind, it examines insurance as an instrument of health promotion policy for improving population health and reducing healthcare costs. The following questions guide this study: (i) How has insurance coverage of employee healthcare in these two countries evolved and changed over time, and why? (ii) What factors incentivize payers (employers and insurers) to incorporate health promotion into their health plans, even if health promotion does not fall within the traditional ambit of insurance? and (iii) How does health promotion through the insurance vehicle operate cost-efficiently and within the Global Health Promotion framework of the World Health Organization (WHO)? US and Philippine public-use datasets were gathered and disaggregated to address these questions. The Affordable Care Act (ACA) in the US and universal healthcare legislation leading to the Universal Healthcare Act (UHC) in the Philippines initiated major paradigmatic shifts in insurance coverage. Health literacy, behavioral change and social equity are objectives pursued in both countries by mandating and/or nudging payers to offer health screenings, wellness and fitness programs and primary preventive and diagnostic services. By providing a means to finance access to promotive health and encourage individual responsibility for risk prevention ('healthism'), insurance influences social attitudes about health. However, our findings indicate that outcomes and relative success vary depending on a country's insurance environment, its regulation and the economic, political and socio-cultural forces that affect or shape it (e.g. private markets vs. multi-payer system). Contextual realities also drive incentives and rewards to payers and users of promotive health benefits against the backdrop of financial risk or exposure.

Schools as Drivers of Capitalist Accumulation Conditional Socialized Reproduction in Shenzhen.

Based on fieldwork in an urbanized village of Shenzhen, this paper analyzes the place of schools in the reproduction of Chinese state capitalism. It retraces the circuit of socialized capital that allows for the social reproduction of the native elite and the exclusion of many migrant workers in the context of Shenzhen's development as a special economic zone and its efforts to upgrade the economy. The native villagers, now forming an urban upper-class of rentiers, have capitalized on their overseas connections and capital accumulation to finance their school, allowing for their elite's upward social mobility after, but also already under Mao. After China's transition to capitalism, this school has served as an asset in generating value in the context of redevelopment and the real estate-driven upgrading of Shenzhen's economy. Property ownership is now a major criterion in points-based systems for accessing school places. I make two interrelated arguments. First, there is a closer relationship between the secondary circuit of socialized capital and the larger circuit of capital than what the literature on social reproduction implies. Second, the conditionality of quality education upon value generation amounts to separating the population deemed worthy of socialized reproduction and the surplus population that is left out. The paper connects diverse strands of social reproduction theory, Althusser's interpellation and ideological state apparatuses, feminist agentive social reproduction theories, and Bourdieu's capital conversion recuperated within a Marxian framework, to provide an integrated approach to social reproduction within capitalism.

Siderophores drive invasion dynamics in bacterial communities through their dual role as public good versus public bad.

Microbial invasions can compromise ecosystem services and spur dysbiosis and disease in hosts. Nevertheless, the mechanisms determining invasion outcomes often remain unclear. Here, we examine the role of iron-scavenging siderophores in driving invasions of Pseudomonas aeruginosa into resident communities of environmental pseudomonads. Siderophores can be 'public goods' by delivering iron to individuals possessing matching receptors; but they can also be 'public bads' by withholding iron from competitors lacking these receptors. Accordingly, siderophores should either promote or impede invasion, depending on their effects on invader and resident growth. Using supernatant feeding and invasion assays, we show that invasion success indeed increased when the invader could use its siderophores to inhibit (public bad) rather than stimulate (public good) resident growth. Conversely, invasion success decreased the more the invader was inhibited by the residents' siderophores. Our findings identify siderophores as a major driver of invasion dynamics in bacterial communities under iron-limited conditions.

Social interaction can select for reduced ability.

Animals, including humans, differ in a wide range of physical and cognitive abilities ranging from measures of running speed and physical strength to learning ability and intelligence. We consider the evolution of ability when individuals interact pairwise over their contribution to a common good. In this interaction, the contribution of each is assumed to be the best given their own ability and the contribution of their partner. Since there is a tendency for individuals to partially compensate for a low contribution by their partner, low-ability individuals can do well. As a consequence, for benefit and cost structures for which individuals have a strong response to partner's contribution, there can be selection for reduced ability. Furthermore, there can be disruptive selection on ability, leading to a bimodal distribution of ability under some modes of inheritance.

The Colorful World of Extracellular Electron Shuttles.

Descriptions of the changeable, striking colors associated with secreted natural products date back well over a century. These molecules can serve as extracellular electron shuttles (EESs) that permit microbes to access substrates at a distance. In this review, we argue that the colorful world of EESs has been too long neglected. Rather than simply serving as a diagnostic attribute of a particular microbial strain, redox-active natural products likely play fundamental, underappreciated roles in the biology of their producers, particularly those that inhabit biofilms. Here, we describe the chemical diversity and potential distribution of EES producers and users, discuss the costs associated with their biosynthesis, and critically evaluate strategies for their economical usage. We hope this review will inspire efforts to identify and explore the importance of EES cycling by a wide range of microorganisms so that their contributions to shaping microbial communities can be better assessed and exploited.