Emery, N. J. (2000). The eyes have it: the neuroethology, function and evolution of social gaze. Neurosci Biobehav Rev, 24(6), 581–604.
Abstract: Gaze is an important component of social interaction. The function, evolution and neurobiology of gaze processing are therefore of interest to a number of researchers. This review discusses the evolutionary role of social gaze in vertebrates (focusing on primates), and a hypothesis that this role has changed substantially for primates compared to other animals. This change may have been driven by morphological changes to the face and eyes of primates, limitations in the facial anatomy of other vertebrates, changes in the ecology of the environment in which primates live, and a necessity to communicate information about the environment, emotional and mental states. The eyes represent different levels of signal value depending on the status, disposition and emotional state of the sender and receiver of such signals. There are regions in the monkey and human brain which contain neurons that respond selectively to faces, bodies and eye gaze. The ability to follow another individual's gaze direction is affected in individuals with autism and other psychopathological disorders, and after particular localized brain lesions. The hypothesis that gaze following is “hard-wired” in the brain, and may be localized within a circuit linking the superior temporal sulcus, amygdala and orbitofrontal cortex is discussed.
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Emery, N. J., & Clayton, N. S. (2004). The Mentality of Crows: Convergent Evolution of Intelligence in Corvids and Apes. Science, 306(5703), 1903–1907.
Abstract: Discussions of the evolution of intelligence have focused on monkeys and apes because of their close evolutionary relationship to humans. Other large-brained social animals, such as corvids, also understand their physical and social worlds. Here we review recent studies of tool manufacture, mental time travel, and social cognition in corvids, and suggest that complex cognition depends on a “tool kit” consisting of causal reasoning, flexibility, imagination, and prospection. Because corvids and apes share these cognitive tools, we argue that complex cognitive abilities evolved multiple times in distantly related species with vastly different brain structures in order to solve similar socioecological problems.
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Emery, N. J., Clayton, N. S., & Frith, C. D. (2007). Introduction. Social intelligence: from brain to culture. Philos Trans R Soc B, 362.
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Emery, N. J., Dally, J. M., & Clayton, N. S. (2004). Western scrub-jays ( Aphelocoma californica) use cognitive strategies to protect their caches from thieving conspecifics. Anim. Cogn., 7(1), 37–43.
Abstract: Food caching birds hide food and recover the caches when supplies are less abundant. There is, however, a risk to this strategy because the caches are susceptible to pilfering by others. Corvids use a number of different strategies to reduce possible cache theft. Scrub-jays with previous experience of pilfering other's caches cached worms in two visuospatially distinct caching trays either in private or in the presence of a conspecific. When these storers had cached in private, they subsequently observed both trays out of reach of a conspecific. When these storers had cached in the presence of a conspecific, they subsequently watched the observer pilfering from one of the trays while the other tray was placed in full view, but out of reach. The storers were then allowed to recover the remaining caches 3 h later. Jays cached more worms when they were observed during caching. At the time of recovery, they re-cached more than if they had cached in private, selectively re-caching outside of the trays in sites unbeknown to potential thieves. In addition, after a single pilfering trial, the jays switched their recovery strategy from predominantly checking their caches (i.e. returning to a cache site to see whether the food remained there) to predominantly eating them. Re-caching remained constant across the three trials. These results suggest that scrub-jays use flexible, cognitive caching and recovery strategies to aid in reducing potential future pilfering of caches by conspecifics.
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Emery, N. J., Seed, A. M., von Bayern, A. M. P., & Clayton, N. S. (2007). Cognitive adaptations of social bonding in birds. Phil. Trans. Biol. Sci., 362(1480), 489–505.
Abstract: The “social intelligence hypothesis” was originally conceived to explain how primates may have evolved their superior intellect and large brains when compared with other animals. Although some birds such as corvids may be intellectually comparable to apes, the same relationship between sociality and brain size seen in primates has not been found for birds, possibly suggesting a role for other non-social factors. But bird sociality is different from primate sociality. Most monkeys and apes form stable groups, whereas most birds are monogamous, and only form large flocks outside of the breeding season. Some birds form lifelong pair bonds and these species tend to have the largest brains relative to body size. Some of these species are known for their intellectual abilities (e.g. corvids and parrots), while others are not (e.g. geese and albatrosses). Although socio-ecological factors may explain some of the differences in brain size and intelligence between corvids/parrots and geese/albatrosses, we predict that the type and quality of the bonded relationship is also critical. Indeed, we present empirical evidence that rook and jackdaw partnerships resemble primate and dolphin alliances. Although social interactions within a pair may seem simple on the surface, we argue that cognition may play an important role in the maintenance of long-term relationships, something we name as “relationship intelligence”.
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MacLean, E., Matthews, L., Hare, B., Nunn, C., Anderson, R., Aureli, F., et al. (2012). How does cognition evolve? Phylogenetic comparative psychology. Anim. Cogn., 15(2), 223–238.
Abstract: Now more than ever animal studies have the potential to test hypotheses regarding how cognition evolves. Comparative psychologists have developed new techniques to probe the cognitive mechanisms underlying animal behavior, and they have become increasingly skillful at adapting methodologies to test multiple species. Meanwhile, evolutionary biologists have generated quantitative approaches to investigate the phylogenetic distribution and function of phenotypic traits, including cognition. In particular, phylogenetic methods can quantitatively (1) test whether specific cognitive abilities are correlated with life history (e.g., lifespan), morphology (e.g., brain size), or socio-ecological variables (e.g., social system), (2) measure how strongly phylogenetic relatedness predicts the distribution of cognitive skills across species, and (3) estimate the ancestral state of a given cognitive trait using measures of cognitive performance from extant species. Phylogenetic methods can also be used to guide the selection of species comparisons that offer the strongest tests of a priori predictions of cognitive evolutionary hypotheses (i.e., phylogenetic targeting). Here, we explain how an integration of comparative psychology and evolutionary biology will answer a host of questions regarding the phylogenetic distribution and history of cognitive traits, as well as the evolutionary processes that drove their evolution.
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Seed, A. M., Clayton, N. S., & Emery, N. J. (2007). Postconflict third-party affiliation in rooks, Corvus frugilegus. Curr Biol, 17(2), 152–158.
Abstract: Conflict features in the lives of many animal species and induces social stress mediated by glucocorticoid hormones [1]. Postconflict affiliation, between former opponents (reconciliation) or between former opponents and a bystander (third-party affiliation), has been suggested as a behavioral mechanism for reducing such stress [2], but has been studied almost exclusively in primates [3]. As with many primates, several bird species live in social groups and form affiliative relationships [4]. Do these distantly related animals also use affiliative behavior to offset the costs of conflict? We studied postconflict affiliation in a captive group of rooks. Unlike polygamous primates, monogamous rooks did not reconcile with former opponents. However, we found clear evidence of third-party affiliation after conflicts. Both initiators and targets of aggression engaged in third-party affiliation with a social partner and employed a specific behavior, bill twining, during the postconflict period. Both former aggressors and uninvolved third parties initiated affiliative contacts. Despite the long history of evolutionary divergence, the pattern of third-party affiliation in rooks is strikingly similar to that observed in tolerant primate species. Furthermore, the absence of reconciliation in rooks makes sense in light of the species differences in social systems.
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Tebbich, S., Seed, A. M., Emery, N. J., & Clayton, N. S. (2007). Non-tool-using rooks, Corvus frugilegus, solve the trap-tube problem. Anim. Cogn., 10(2), 225–231.
Abstract: The trap-tube problem is used to assess whether an individual is able to foresee the outcome of its actions. To solve the task, an animal must use a tool to push a piece of food out of a tube, which has a trap along its length. An animal may learn to avoid the trap through a rule based on associative processes, e.g. using the distance of trap or food as a cue, or by understanding relations between cause and effect. This task has been used to test physical cognition in a number of tool-using species, but never a non-tool-user. We developed an experimental design that enabled us to test non-tool-using rooks, Corvus frugilegus. Our modification of the task removed the cognitive requirements of active tool use but still allowed us to test whether rooks can solve the trap-tube problem, and if so how. Additionally, we developed two new control tasks to determine whether rooks were able to transfer knowledge to similar, but novel problems, thus revealing more about the mechanisms involved in solving the task. We found that three out of seven rooks solved the modified trap-tube problem task, showing that the ability to solve the trap-tube problem is not restricted to tool-using animals. We found no evidence that the birds solved the task using an understanding of its causal properties, given that none of the birds passed the novel transfer tasks.
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Van Horik, J., Clayton, N., & Emery, N. (2012). Convergent evolution of cognition in Corvids, Apes and other animals. In J. Vonk, & T. Shackelford (Eds.), Oxford Handbook of Comparative Evolutionary Psychology. New York: Oxford University Press.
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Van Horik, J., Clayton, N., & Emery, N. Oxford Handbook of Comparative Evolutionary Psychology (J. Vonk, & T. Shackelford, Eds.). New York: Oxford University Press.
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