Super-additive cooperation.

Repeated interactions provide an evolutionary explanation for one-shot human cooperation that is counterintuitive but orthodox1-3. Intergroup competition4-7 provides an explanation that is intuitive but heterodox. Here, using models and a behavioural experiment, we show that neither mechanism reliably supports cooperation. Ambiguous reciprocity, a class of strategies that is generally ignored in models of reciprocal altruism, undermines cooperation under repeated interactions. This finding challenges repeated interactions as an evolutionary explanation for cooperation in general, which further challenges the claim that repeated interactions in the past can explain one-shot cooperation in the present. Intergroup competitions also do not reliably support cooperation because groups quickly become extremely similar, which limits scope for group selection. Moreover, even if groups vary, group competitions may generate little group selection for multiple reasons. Cooperative groups, for example, may tend to compete against each other8. Whereas repeated interactions and group competitions do not support cooperation by themselves, combining them triggers powerful synergies because group competitions constrain the corrosive effect of ambiguous reciprocity. Evolved strategies often consist of cooperative reciprocity with ingroup partners and uncooperative reciprocity with outgroup partners. Results from a behavioural experiment in Papua New Guinea fit exactly this pattern. They thus suggest neither an evolutionary history of repeated interactions without group competition nor a history of group competition without repeated interactions. Instead, our results suggest social motives that evolved under the joint influence of both mechanisms.

Dynamic behaviour restructuring mediates dopamine-dependent credit assignment.

Animals exhibit a diverse behavioural repertoire when exploring new environments and can learn which actions or action sequences produce positive outcomes. Dopamine release after encountering a reward is critical for reinforcing reward-producing actions1-3. However, it has been challenging to understand how credit is assigned to the exact action that produced the dopamine release during continuous behaviour. Here we investigated this problem in mice using a self-stimulation paradigm in which specific spontaneous movements triggered optogenetic stimulation of dopaminergic neurons. Dopamine self-stimulation rapidly and dynamically changes the structure of the entire behavioural repertoire. Initial stimulations reinforced not only the stimulation-producing target action, but also actions similar to the target action and actions that occurred a few seconds before stimulation. Repeated pairings led to a gradual refinement of the behavioural repertoire to home in on the target action. Reinforcement of action sequences revealed further temporal dependencies of refinement. Action pairs spontaneously separated by long time intervals promoted a stepwise credit assignment, with early refinement of actions most proximal to stimulation and subsequent refinement of more distal actions. Thus, a retrospective reinforcement mechanism promotes not only reinforcement, but also gradual refinement of the entire behavioural repertoire to assign credit to specific actions and action sequences that lead to dopamine release.

Expertise increases planning depth in human gameplay.

A hallmark of human intelligence is the ability to plan multiple steps into the future1,2. Despite decades of research3-5, it is still debated whether skilled decision-makers plan more steps ahead than novices6-8. Traditionally, the study of expertise in planning has used board games such as chess, but the complexity of these games poses a barrier to quantitative estimates of planning depth. Conversely, common planning tasks in cognitive science often have a lower complexity9,10 and impose a ceiling for the depth to which any player can plan. Here we investigate expertise in a complex board game that offers ample opportunity for skilled players to plan deeply. We use model fitting methods to show that human behaviour can be captured using a computational cognitive model based on heuristic search. To validate this model, we predict human choices, response times and eye movements. We also perform a Turing test and a reconstruction experiment. Using the model, we find robust evidence for increased planning depth with expertise in both laboratory and large-scale mobile data. Experts memorize and reconstruct board features more accurately. Using complex tasks combined with precise behavioural modelling might expand our understanding of human planning and help to bridge the gap with progress in artificial intelligence.

People construct simplified mental representations to plan.

One of the most striking features of human cognition is the ability to plan. Two aspects of human planning stand out-its efficiency and flexibility. Efficiency is especially impressive because plans must often be made in complex environments, and yet people successfully plan solutions to many everyday problems despite having limited cognitive resources1-3. Standard accounts in psychology, economics and artificial intelligence have suggested that human planning succeeds because people have a complete representation of a task and then use heuristics to plan future actions in that representation4-11. However, this approach generally assumes that task representations are fixed. Here we propose that task representations can be controlled and that such control provides opportunities to quickly simplify problems and more easily reason about them. We propose a computational account of this simplification process and, in a series of preregistered behavioural experiments, show that it is subject to online cognitive control12-14 and that people optimally balance the complexity of a task representation and its utility for planning and acting. These results demonstrate how strategically perceiving and conceiving problems facilitates the effective use of limited cognitive resources.

People systematically overlook subtractive changes.

Improving objects, ideas or situations-whether a designer seeks to advance technology, a writer seeks to strengthen an argument or a manager seeks to encourage desired behaviour-requires a mental search for possible changes1-3. We investigated whether people are as likely to consider changes that subtract components from an object, idea or situation as they are to consider changes that add new components. People typically consider a limited number of promising ideas in order to manage the cognitive burden of searching through all possible ideas, but this can lead them to accept adequate solutions without considering potentially superior alternatives4-10. Here we show that people systematically default to searching for additive transformations, and consequently overlook subtractive transformations. Across eight experiments, participants were less likely to identify advantageous subtractive changes when the task did not (versus did) cue them to consider subtraction, when they had only one opportunity (versus several) to recognize the shortcomings of an additive search strategy or when they were under a higher (versus lower) cognitive load. Defaulting to searches for additive changes may be one reason that people struggle to mitigate overburdened schedules11, institutional red tape12 and damaging effects on the planet13,14.

Indirect reciprocity undermines indirect reciprocity destabilizing large-scale cooperation.

Previous models suggest that indirect reciprocity (reputation) can stabilize large-scale human cooperation [K. Panchanathan, R. Boyd, Nature 432, 499-502 (2004)]. The logic behind these models and experiments [J. Gross et al., Sci. Adv. 9, eadd8289 (2023) and O. P. Hauser, A. Hendriks, D. G. Rand, M. A. Nowak, Sci. Rep. 6, 36079 (2016)] is that a strategy in which individuals conditionally aid others based on their reputation for engaging in costly cooperative behavior serves as a punishment that incentivizes large-scale cooperation without the second-order free-rider problem. However, these models and experiments fail to account for individuals belonging to multiple groups with reputations that can be in conflict. Here, we extend these models such that individuals belong to a smaller, "local" group embedded within a larger, "global" group. This introduces competing strategies for conditionally aiding others based on their cooperative behavior in the local or global group. Our analyses reveal that the reputation for cooperation in the smaller local group can undermine cooperation in the larger global group, even when the theoretical maximum payoffs are higher in the larger global group. This model reveals that indirect reciprocity alone is insufficient for stabilizing large-scale human cooperation because cooperation at one scale can be considered defection at another. These results deepen the puzzle of large-scale human cooperation.

Two views on the cognitive brain.

Cognition can be defined as computation over meaningful representations in the brain to produce adaptive behaviour. There are two views on the relationship between cognition and the brain that are largely implicit in the literature. The Sherringtonian view seeks to explain cognition as the result of operations on signals performed at nodes in a network and passed between them that are implemented by specific neurons and their connections in circuits in the brain. The contrasting Hopfieldian view explains cognition as the result of transformations between or movement within representational spaces that are implemented by neural populations. Thus, the Hopfieldian view relegates details regarding the identity of and connections between specific neurons to the status of secondary explainers. Only the Hopfieldian approach has the representational and computational resources needed to develop novel neurofunctional objects that can serve as primary explainers of cognition.

Evolutionary stability of cooperation in indirect reciprocity under noisy and private assessment.

Indirect reciprocity is a mechanism that explains large-scale cooperation in humans. In indirect reciprocity, individuals use reputations to choose whether or not to cooperate with a partner and update others' reputations. A major question is how the rules to choose their actions and the rules to update reputations evolve. In the public reputation case where all individuals share the evaluation of others, social norms called Simple Standing (SS) and Stern Judging (SJ) have been known to maintain cooperation. However, in the case of private assessment where individuals independently evaluate others, the mechanism of maintenance of cooperation is still largely unknown. This study theoretically shows for the first time that cooperation by indirect reciprocity can be evolutionarily stable under private assessment. Specifically, we find that SS can be stable, but SJ can never be. This is intuitive because SS can correct interpersonal discrepancies in reputations through its simplicity. On the other hand, SJ is too complicated to avoid an accumulation of errors, which leads to the collapse of cooperation. We conclude that moderate simplicity is a key to stable cooperation under the private assessment. Our result provides a theoretical basis for the evolution of human cooperation.

Emergence of specialized third-party enforcement.

The question of how cooperation evolves and is maintained among nonkin is central to the biological, social, and behavioral sciences. Previous research has focused on explaining how cooperation in social dilemmas can be maintained by direct and indirect reciprocity among the participants of the social dilemma. However, in complex human societies, both modern and ancient, cooperation is frequently maintained by means of specialized third-party enforcement. We provide an evolutionary-game-theoretic model that explains how specialized third-party enforcement of cooperation (specialized reciprocity) can emerge. A population consists of producers and enforcers. First, producers engage in a joint undertaking represented by a prisoner's dilemma. They are paired randomly and receive no information about their partner's history, which precludes direct and indirect reciprocity. Then, enforcers tax producers and may punish their clients. Finally, the enforcers are randomly paired and may try to grab resources from each other. In order to sustain producer cooperation, enforcers must punish defecting producers, but punishing is costly to enforcers. We show that the threat of potential intraenforcer conflict can incentivize enforcers to engage in costly punishment of producers, provided they are sufficiently informed to maintain a reputation system. That is, the "guards" are guarded by the guards themselves. We demonstrate the key mechanisms analytically and corroborate our results with numerical simulations.

Evolution of norms for judging social behavior.

Reputations provide a powerful mechanism to sustain cooperation, as individuals cooperate with those of good social standing. But how should someone's reputation be updated as we observe their social behavior, and when will a population converge on a shared norm for judging behavior? Here, we develop a mathematical model of cooperation conditioned on reputations, for a population that is stratified into groups. Each group may subscribe to a different social norm for assessing reputations and so norms compete as individuals choose to move from one group to another. We show that a group initially comprising a minority of the population may nonetheless overtake the entire population-especially if it adopts the Stern Judging norm, which assigns a bad reputation to individuals who cooperate with those of bad standing. When individuals do not change group membership, stratifying reputation information into groups tends to destabilize cooperation, unless individuals are strongly insular and favor in-group social interactions. We discuss the implications of our results for the structure of information flow in a population and for the evolution of social norms of judgment.

Emergence of two distinct spatial folds in a pair of plant virus proteins encoded by nested genes.

Virus genomes may encode overlapping or nested open reading frames that increase their coding capacity. It is not known whether the constraints on spatial structures of the two encoded proteins limit the evolvability of nested genes. We examine the evolution of a pair of proteins, p22 and p19, encoded by nested genes in plant viruses from the genus Tombusvirus. The known structure of p19, a suppressor of RNA silencing, belongs to the RAGNYA fold from the alpha+beta class. The structure of p22, the cell-to-cell movement protein from the 30K family widespread in plant viruses, is predicted with the AlphaFold approach, suggesting a single jelly-roll fold core from the all-beta class, structurally similar to capsid proteins from plant and animal viruses. The nucleotide and codon preferences impose modest constraints on the types of secondary structures encoded in the alternative reading frames, nonetheless allowing for compact, well-ordered folds from different structural classes in two similarly-sized nested proteins. Tombusvirus p22 emerged through radiation of the widespread 30K family, which evolved by duplication of a virus capsid protein early in the evolution of plant viruses, whereas lineage-specific p19 may have emerged by a stepwise increase in the length of the overprinted gene and incremental acquisition of functionally active secondary structure elements by the protein product. This evolution of p19 toward the RAGNYA fold represents one of the first documented examples of protein structure convergence in naturally occurring proteins.

When do stereotypes undermine indirect reciprocity?

Social reputations provide a powerful mechanism to stimulate human cooperation, but observing individual reputations can be cognitively costly. To ease this burden, people may rely on proxies such as stereotypes, or generalized reputations assigned to groups. Such stereotypes are less accurate than individual reputations, and so they could disrupt the positive feedback between altruistic behavior and social standing, undermining cooperation. How do stereotypes impact cooperation by indirect reciprocity? We develop a theoretical model of group-structured populations in which individuals are assigned either individual reputations based on their own actions or stereotyped reputations based on their groups' behavior. We find that using stereotypes can produce either more or less cooperation than using individual reputations, depending on how widely reputations are shared. Deleterious outcomes can arise when individuals adapt their propensity to stereotype. Stereotyping behavior can spread and can be difficult to displace, even when it compromises collective cooperation and even though it makes a population vulnerable to invasion by defectors. We discuss the implications of our results for the prevalence of stereotyping and for reputation-based cooperation in structured populations.

Human decision making balances reward maximization and policy compression.

Policy compression is a computational framework that describes how capacity-limited agents trade reward for simpler action policies to reduce cognitive cost. In this study, we present behavioral evidence that humans prefer simpler policies, as predicted by a capacity-limited reinforcement learning model. Across a set of tasks, we find that people exploit structure in the relationships between states, actions, and rewards to "compress" their policies. In particular, compressed policies are systematically biased towards actions with high marginal probability, thereby discarding some state information. This bias is greater when there is redundancy in the reward-maximizing action policy across states, and increases with memory load. These results could not be explained qualitatively or quantitatively by models that did not make use of policy compression under a capacity limit. We also confirmed the prediction that time pressure should further reduce policy complexity and increase action bias, based on the hypothesis that actions are selected via time-dependent decoding of a compressed code. These findings contribute to a deeper understanding of how humans adapt their decision-making strategies under cognitive resource constraints.

Beliefs, compulsive behavior and reduced confidence in control.

OCD has been conceptualized as a disorder arising from dysfunctional beliefs, such as overestimating threats or pathological doubts. Yet, how these beliefs lead to compulsions and obsessions remains unclear. Here, we develop a computational model to examine the specific beliefs that trigger and sustain compulsive behavior in a simple symptom-provoking scenario. Our results demonstrate that a single belief disturbance-a lack of confidence in the effectiveness of one's preventive (harm-avoiding) actions-can trigger and maintain compulsions and is directly linked to compulsion severity. This distrust can further explain a number of seemingly unrelated phenomena in OCD, including the role of not-just-right feelings, the link to intolerance to uncertainty, perfectionism, and overestimation of threat, and deficits in reversal and state learning. Our simulations shed new light on which underlying beliefs drive compulsive behavior and highlight the important role of perceived ability to exert control for OCD.

A low-dimensional approximation of optimal confidence.

Human decision making is accompanied by a sense of confidence. According to Bayesian decision theory, confidence reflects the learned probability of making a correct response, given available data (e.g., accumulated stimulus evidence and response time). Although optimal, independently learning these probabilities for all possible data combinations is computationally intractable. Here, we describe a novel model of confidence implementing a low-dimensional approximation of this optimal yet intractable solution. This model allows efficient estimation of confidence, while at the same time accounting for idiosyncrasies, different kinds of biases and deviation from the optimal probability correct. Our model dissociates confidence biases resulting from the estimate of the reliability of evidence by individuals (captured by parameter α), from confidence biases resulting from general stimulus independent under and overconfidence (captured by parameter ß). We provide empirical evidence that this model accurately fits both choice data (accuracy, response time) and trial-by-trial confidence ratings simultaneously. Finally, we test and empirically validate two novel predictions of the model, namely that 1) changes in confidence can be independent of performance and 2) selectively manipulating each parameter of our model leads to distinct patterns of confidence judgments. As a tractable and flexible account of the computation of confidence, our model offers a clear framework to interpret and further resolve different forms of confidence biases.

Joint modeling of choices and reaction times based on Bayesian contextual behavioral control.

In cognitive neuroscience and psychology, reaction times are an important behavioral measure. However, in instrumental learning and goal-directed decision making experiments, findings often rely only on choice probabilities from a value-based model, instead of reaction times. Recent advancements have shown that it is possible to connect value-based decision models with reaction time models. However, typically these models do not provide an integrated account of both value-based choices and reaction times, but simply link two types of models. Here, we propose a novel integrative joint model of both choices and reaction times by combining a computational account of Bayesian sequential decision making with a sampling procedure. This allows us to describe how internal uncertainty in the planning process shapes reaction time distributions. Specifically, we use a recent context-specific Bayesian forward planning model which we extend by a Markov chain Monte Carlo (MCMC) sampler to obtain both choices and reaction times. As we will show this makes the sampler an integral part of the decision making process and enables us to reproduce, using simulations, well-known experimental findings in value based-decision making as well as classical inhibition and switching tasks. Specifically, we use the proposed model to explain both choice behavior and reaction times in instrumental learning and automatized behavior, in the Eriksen flanker task and in task switching. These findings show that the proposed joint behavioral model may describe common underlying processes in these different decision making paradigms.

Psychological Flexibility, Chronic Pain, and Health.

Psychological flexibility is a model of human performance and well-being. It essentially entails an approach to life circumstances that includes openness, awareness, and engagement. It has roots in behavior analysis, and it is linked to a philosophy of science called functional contextualism and to a specific therapy approach called Acceptance and Commitment Therapy. One of the earliest and most developed research areas in which this model and therapy have been applied is chronic pain. This review describes psychological flexibility and its facets in more detail, sets them in a context of relevant psychological models, and examines related assessment and treatment methods. It also examines evidence, current challenges, and future directions. It is proposed that psychological flexibility, or an expanded model very much like it, could provide a basis for integrating current research and treatment approaches in chronic pain and health generally. This, in turn, could produce improved treatments for people with chronic pain and other conditions.

Individuals with adverse childhood experiences explore less and underweight reward feedback.

Adverse childhood experiences (ACEs) are extreme stressors that lead to negative psychosocial outcomes in adulthood. Nonhuman animals explore less after exposure to early stress. Therefore, in this preregistered study, we hypothesized that reduced exploration following ACEs would also be evident in human adults. Further, we predicted that adults with ACEs, in a foraging task, would adopt a decision-making policy that relies on the most-recent reward feedback, a rational strategy for unstable environments. We analyzed data from 145 adult participants, 47 with four or more ACEs and 98 with fewer than four ACEs. In the foraging task, participants evaluated the trade-off between exploiting a known patch with diminishing rewards and exploring a novel one with a fresh distribution of rewards. Using computational modeling, we quantified the degree to which participants' decisions weighted recent feedback. As predicted, participants with ACEs explored less. However, contrary to our hypothesis, they underweighted recent feedback. These unexpected findings indicate that early adversity may dampen reward sensitivity. Our results may help to identify cognitive mechanisms that link childhood trauma to the onset of psychopathology.

Optimizing the human learnability of abstract network representations.

Precisely how humans process relational patterns of information in knowledge, language, music, and society is not well understood. Prior work in the field of statistical learning has demonstrated that humans process such information by building internal models of the underlying network structure. However, these mental maps are often inaccurate due to limitations in human information processing. The existence of such limitations raises clear questions: Given a target network that one wishes for a human to learn, what network should one present to the human? Should one simply present the target network as-is, or should one emphasize certain parts of the network to proactively mitigate expected errors in learning? To investigate these questions, we study the optimization of network learnability in a computational model of human learning. Evaluating an array of synthetic and real-world networks, we find that learnability is enhanced by reinforcing connections within modules or clusters. In contrast, when networks contain significant core-periphery structure, we find that learnability is best optimized by reinforcing peripheral edges between low-degree nodes. Overall, our findings suggest that the accuracy of human network learning can be systematically enhanced by targeted emphasis and de-emphasis of prescribed sectors of information.

Americans discount the effect of friction on voter turnout.

Whether or not someone turns out to vote depends on their beliefs (such as partisanship or sense of civic duty) and on friction-external barriers such as long travel distance to the polls. In this exploratory study, we tested whether people underestimate the effect of friction on turnout and overestimate the effect of beliefs. We surveyed a representative sample of eligible US voters before and after the 2020 election (n = 1,280). Participants' perceptions consistently underemphasized friction and overemphasized beliefs (mean d = 0.94). In participants' open-text explanations, 91% of participants listed beliefs, compared with just 12% that listed friction. In contrast, turnout was shaped by beliefs only slightly more than friction. The actual belief-friction difference was about one-fourth the size of participants' perceptions (d = 0.24). This bias emerged across a range of survey measures (open- and close-ended; other- and self-judgments) and was implicated in downstream consequences such as support for friction-imposing policies and failing to plan one's vote.