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WIREs Cogn Sci
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Primate empathy: three factors and their combinations for empathy‐related phenomena

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Empathy as a research topic is receiving increasing attention, although there seems some confusion on the definition of empathy across different fields. Frans de Waal (de Waal FBM. Putting the altruism back into altruism: the evolution of empathy. Annu Rev Psychol 2008, 59:279–300. doi:10.1146/annurev.psych.59.103006.093625) used empathy as an umbrella term and proposed a comprehensive model for the evolution of empathy with some of its basic elements in nonhuman animals. In de Waal's model, empathy consists of several layers distinguished by required cognitive levels; the perception‐action mechanism plays the core role for connecting ourself and others. Then, human‐like empathy such as perspective‐taking develops in outer layers according to cognitive sophistication, leading to prosocial acts such as targeted helping. I agree that animals demonstrate many empathy‐related phenomena; however, the species differences and the level of cognitive sophistication of the phenomena might be interpreted in another way than this simple linearly developing model. Our recent studies with chimpanzees showed that their perspective‐taking ability does not necessarily lead to proactive helping behavior. Herein, as a springboard for further studies, I reorganize the empathy‐related phenomena by proposing a combination model instead of the linear development model. This combination model is composed of three organizing factors: matching with others, understanding of others, and prosociality. With these three factors and their combinations, most empathy‐related matters can be categorized and mapped to appropriate context; this may be a good first step to discuss the evolution of empathy in relation to the neural connections in human and nonhuman animal brains. I would like to propose further comparative studies, especially from the viewpoint of Homo‐Pan (chimpanzee and bonobo) comparison. WIREs Cogn Sci 2017, 8:e1431. doi: 10.1002/wcs.1431 This article is categorized under: Cognitive Biology > Evolutionary Roots of Cognition Psychology > Comparative Psychology Neuroscience > Cognition
Chimpanzees' flexible targeted helping upon request. The experiments were conducted for two conditions: ‘can see’ conditions (a) and (c), in which a helper chimpanzee could see the partner's tool‐use (‘stick’ or ‘straw’), and ‘cannot see’ condition (b), in which the partition was opaque so that a helper could not see the partner's situation, although the partner could demonstrate a request through the partition's small opening. The graphs represent helpers’ first tool selection from seven objects that was offered to their conspecific partner. This shows that chimpanzees can understand what specific tool the partner needs when they see the partner's situation although this understanding does not provoke their proactive helping. (Reprinted with permission from Ref . Copyright 2012 Proceedings of the National Academy of Sciences USA)
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Bonobos’ fruit sharing. They often share daily fruit items that are much more easily available than meat, and this might strengthen their social bond. The bonobo is one of the most interesting and important species in which to investigate the evolution of empathy from comparative cognitive viewpoints. (Photograph by Shinya Yamamoto at Wamba, the Democratic Republic of the Congo.)
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Three major facets and their combinations for empathy‐related phenomena. Any one of the empathy‐related phenomena can fall into one of seven categories (a‐g). Recent studies have revealed evidence, or at least some elements, for each category in nonhuman animals, but the range of animal species that demonstrate the phenomena depends on the categories. It seems that the combination of two or more factors in this model requires considerable evolutionary development. See the text for details.
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Neuroscience > Cognition
Cognitive Biology > Evolutionary Roots of Cognition
Psychology > Comparative Psychology

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