Eleven predictions derived from the recalibrational theory of anger were tested. This theory proposes that anger is produced by a neurocognitive program engineered by natural selection to use bargaining tactics to resolve conflicts of interest in favor of the angry individual. The program is designed to orchestrate two interpersonal negotiating tactics (conditionally inflicting costs or conditionally withholding benefits) to incentivize the target of the anger to place greater weight on the welfare of the angry individual. Individuals with enhanced abilities to inflict costs (e.g., stronger individuals) or to confer benefits (e.g., attractive individuals) have a better bargaining position in conflicts; hence, it was predicted that such individuals will be more prone to anger, prevail more in conflicts of interest, and consider themselves entitled to better treatment. These predictions were confirmed. Consistent with an evolutionary analysis, the effect of strength on anger was greater for men and the effect of attractiveness on anger was greater for women. Also as predicted, stronger men had a greater history of fighting than weaker men, and more strongly endorsed the efficacy of force to resolve conflicts-both in interpersonal and international conflicts. The fact that stronger men favored greater use of military force in international conflicts provides evidence that the internal logic of the anger program reflects the ancestral payoffs characteristic of a small-scale social world rather than rational assessments of modern payoffs in large populations.aggression ͉ evolutionary psychology ͉ recalibrational theory ͉ welfare tradeoff ratio A nger is part of the basic biology of the human species. It spontaneously appears in infancy (1, 2), is effectively universal in its distribution across cultures and individuals (3, 4), and has a species-typical neural basis (5). To understand the evolutionary biology of anger, however, it is also important to characterize its evolved function-that is, what (if anything) was anger engineered by natural selection to accomplish? A recent model-the recalibrational theory of anger-hypothesizes that the regulatory program governing anger evolved in the service of bargaining, to resolve conflicts of interest in favor of the angry individual (6, 7). More precisely, the function of anger is to orchestrate behavior in the angry individual that creates incentives in the target of the anger to recalibrate upwards the weight he or she puts on the welfare of the angry individual. Here, we report empirical tests of 11 predictions derived from this model. Engineering AnalysisAnger, Welfare Tradeoff Ratios (WTRs), and Bargaining. In social species, actions undertaken by one individual commonly have impacts on the welfare of others (measured in fitness, or in other currencies). Consequently, neurocognitive programs in social species should have been designed by selection to solve the following computational adaptive problem: For a given choice set involving self and other, how much weight shou...

Selection in species with aggressive social interactions favours the evolution of cognitive mechanisms for assessing physical formidability (fighting ability or resource-holding potential). The ability to accurately assess formidability in conspecifics has been documented in a number of non-human species, but has not been demonstrated in humans. Here, we report tests supporting the hypothesis that the human cognitive architecture includes mechanisms that assess fighting ability-mechanisms that focus on correlates of upper-body strength. Across diverse samples of targets that included US college students, Bolivian horticulturalists and Andean pastoralists, subjects in the US were able to accurately estimate the physical strength of male targets from photos of their bodies and faces. Hierarchical linear modelling shows that subjects were extracting cues of strength that were largely independent of height, weight and age, and that corresponded most strongly to objective measures of upper-body strength-even when the face was all that was available for inspection. Estimates of women's strength were less accurate, but still significant. These studies are the first empirical demonstration that, for humans, judgements of strength and judgements of fighting ability not only track each other, but accurately track actual upper-body strength.

Recent research has shown that humans, like many other animals, have a specialization for assessing fighting ability from visual cues. Because it is probable that the voice contains cues of strength and formidability that are not available visually, we predicted that selection has also equipped humans with the ability to estimate physical strength from the voice. We found that subjects accurately assessed upper-body strength in voices taken from eight samples across four distinct populations and language groups: the Tsimane of Bolivia, Andean herder-horticulturalists and United States and Romanian college students. Regardless of whether raters were told to assess height, weight, strength or fighting ability, they produced similar ratings that tracked upper-body strength independent of height and weight. Male voices were more accurately assessed than female voices, which is consistent with ethnographic data showing a greater tendency among males to engage in violent aggression. Raters extracted information about strength from the voice that was not supplied from visual cues, and were accurate with both familiar and unfamiliar languages. These results provide, to our knowledge, the first direct evidence that both men and women can accurately assess men's physical strength from the voice, and suggest that estimates of strength are used to assess fighting ability.

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Fighting ability, although recognized as fundamental to intrasexual competition in many nonhuman species, has received little attention as an explanatory variable in the social sciences. Multiple lines of evidence from archaeology, criminology, anthropology, physiology, and psychology suggest that fighting ability was a crucial aspect of intrasexual competition for ancestral human males, and this has contributed to the evolution of numerous physical and psychological sex differences. Because fighting ability was relevant to many domains of interaction, male psychology should have evolved such that a man's attitudes and behavioral responses are calibrated according to his formidability. Data are reviewed showing that better fighters feel entitled to better outcomes, set lower thresholds for anger/aggression, have self-favoring political attitudes, and believe more in the utility of warfare. New data are presented showing that among Hollywood actors, those selected for their physical strength (i.e., action stars) are more likely to believe in the utility of warfare.

Humans can accurately extract information about men's formidability from their faces; however, the actual facial cues that inform these judgments have not been established. Here, through three studies, we test the hypothesis that bizygomatic width (i.e. facial width-to-height ratio, fWHR) covaries with actual physical formidability (hypothesis #1) and that humans use this cue when making assessments of formidability (hypothesis #2). Our data confirm that fWHR is predictive of actual fighting ability among professional combatants (study 1). We further show that subjects' assessments of formidability covary with the target's fWHR on natural faces (study 2), computer-generated images of strong and weak faces (study 2), and experimentally manipulated computer-generated faces (study 3). These results support the hypothesis that bizygomatic width is a cue of formidability that is assessed during agonistic encounters.

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a b s t r a c tAccording to the recalibrational theory of anger, anger is a computationally complex cognitive system that evolved to bargain for better treatment. Anger coordinates facial expressions, vocal changes, verbal arguments, the withholding of benefits, the deployment of aggression, and a suite of other cognitive and physiological variables in the service of leveraging bargaining position into better outcomes. The prototypical trigger of anger is an indication that the offender places too little weight on the angry individual's welfare when making decisions, i.e. the offender has too low a welfare tradeoff ratio (WTR) toward the angry individual. Twenty-three experiments in six cultures, including a group of foragers in the Ecuadorian Amazon, tested six predictions about the computational structure of anger derived from the recalibrational theory. Subjects judged that anger would intensify when: (i) the cost was large, (ii) the benefit the offender received from imposing the cost was small, or (iii) the offender imposed the cost despite knowing that the angered individual was the person to be harmed. Additionally, anger-based arguments conformed to a conceptual grammar of anger, such that offenders were inclined to argue that they held a high WTR toward the victim, e.g., ''the cost I imposed on you was small", ''the benefit I gained was large", or ''I didn't know it was you I was harming." These results replicated across all six tested cultures: the US, Australia, Turkey, Romania, India, and Shuar hunter-horticulturalists in Ecuador. Results contradict key predictions about anger based on equity theory and social constructivism.