BackgroundThe evolution of altruism has been explained mainly from ultimate perspectives. However, it remains to be investigated from a proximate point of view how and in which situations such social propensity is achieved. We investigated chimpanzees' targeted helping in a tool transfer paradigm, and discuss the similarities and differences in altruism between humans and chimpanzees. Previously it has been suggested that chimpanzees help human experimenters by retrieving an object which the experimenter is trying to reach. In the present study, we investigated the importance of communicative interactions between chimpanzees themselves and the influence of conspecific partner's request on chimpanzees' targeted helping.Methodology/Principal FindingsWe presented two tool-use situations (a stick-use situation and a straw-use situation) in two adjacent booths, and supplied non-corresponding tools to paired chimpanzees in the two booths. For example, a chimpanzee in the stick-use situation was supplied with a straw, and the partner in the straw-use situation possessed a stick. Spontaneous tool transfer was observed between paired chimpanzees. The tool transfer events occurred predominantly following recipients' request. Even without any hope of reciprocation from the partner, the chimpanzees continued to help the partner as long as the partner required help.Conclusions/SignificanceThese results provide further evidence for altruistic helping in chimpanzees in the absence of direct personal gain or even immediate reciprocation. Our findings additionally highlight the importance of request as a proximate mechanism motivating prosocial behavior in chimpanzees whether between kin or non-kin individuals and the possible confounding effect of dominance on the symmetry of such interactions. Finally, in contrast to humans, our study suggests that chimpanzees rarely perform acts of voluntary altruism. Voluntary altruism in chimpanzees is not necessarily prompted by simple observation of another's struggle to attain a goal and therefore an accurate understanding of others' desires in the absence of communicative signals.

Humans extensively help others altruistically, which plays an important role in maintaining cooperative societies. Although some nonhuman animals are also capable of helping others altruistically, humans are considered unique in our voluntary helping and our variety of helping behaviors. Many still believe that this is because only humans can understand others’ goals due to our unique “theory of mind” abilities, especially shared intentionality. However, we know little of the cognitive mechanisms underlying helping in nonhuman animals, especially if and how they understand others’ goals. The present study provides the empirical evidence for flexible targeted helping depending on conspecifics’ needs in chimpanzees. The subjects of this study selected an appropriate tool from a random set of seven objects to transfer to a conspecific partner confronted with differing tool-use situations, indicating that they understood what their partner needed. This targeted helping, (i.e., selecting the appropriate tool to transfer), was observed only when the helpers could visually assess their partner's situation. If visual access was obstructed, the chimpanzees still tried to help their partner upon request, but failed to select and donate the appropriate tool needed by their partner. These results suggest that the limitation in chimpanzees’ voluntary helping is not necessarily due to failure in understanding others’ goals. Chimpanzees can understand conspecifics’ goals and demonstrate cognitively advanced targeted helping as long as they are able to visually evaluate their conspecifics’ predicament. However, they will seldom help others without direct request for help.

BackgroundThe evidence for culture in non-human animals has been growing incrementally over the past two decades. However, the ability for cumulative cultural evolution, with successive generations building on earlier achievements, in non-human animals remains debated. Faithful social learning of incremental improvements in technique is considered to be a defining feature of human culture, differentiating human from non-human cultures. This study presents the first experimental evidence for chimpanzees' social transmission of a more efficient tool-use technique invented by a conspecific group member.Methodology/Principal FindingsThe chimpanzees were provided with a straw-tube, and spontaneously demonstrated two different techniques in obtaining juice through a small hole: “dipping” and “straw-sucking”. Both the “dipping” and “straw-sucking” techniques depended on the use of the same tool (straw-tube) for the same target (juice) accessible from exactly the same location (small hole 1 cm in diameter). Therefore the difference between “dipping” and “straw-sucking” was only in “technique”. Although the two techniques differed significantly in their efficiency, their cognitive and perceptuo-motor complexity were comparable. All five chimpanzees who initially performed the “dipping” technique switched to using the more efficient “straw-sucking” technique upon observing a conspecific or human demonstrate the more proficient alternate “straw-sucking” technique.Conclusions/SignificanceThe social learning mechanism involved here was clearly not local or stimulus enhancement, but imitation or emulation of a tool-use technique. When there is no biologically relevant difference in cognitive or perceptuo-motor complexity between two techniques, and when chimpanzees are dissatisfied with their own technique, chimpanzees may socially learn an improved technique upon close observation of a proficient demonstrator. This study provides important insights into the cognitive basis for cumulative culture in chimpanzees, and also suggests possible conditions in which cumulative cultural evolution could arise even in non-human animals.

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Special thanks are due to Viana do Castelo city for supporting our project. We are also grateful to Agostinho Costinha, the director of Descubra Minho, Lourenço Almada of Associação O Caminho do Garrano. We also thank the villagers in Montaria for their support during our stay, Tetsuro Matsuzawa for the generous guidance throughout the study, and Dora Biro and Valéria Romano for helpful comments on an earlier version of our manuscript. The study was financially supported by grants from the Japan Society for the Promotion of Science (JSPS core-to-core CCSN and JSPS-LGP-U04 to Tetsuro Matsuzawa, KAKENHI Nos. 15H01619 and 15H05309 to Shinya Yamamoto) and the Ministry of Education, Culture, Sports, Science, and Technology in Japan (MEXT No.16H06283 to Tetsuro Matsuzawa). We thank Lilly Gray and Adam Phillips, PhD from Edanz Group (www.edanzediting.com/ac) for editing a draft of this manuscript.

The study of non-human multilevel societies can give us insights into how group-level relationships function and are maintained in a social system, but their mechanisms are still poorly understood. The aim of this study was to apply spatial association data obtained from drones to verify the presence of a multilevel structure in a feral horse society. We took aerial photos of individuals that appeared in pre-fixed areas and collected positional data. The threshold distance of the association was defined based on the distribution pattern of the inter-individual distance. The association rates of individuals showed bimodality, suggesting the presence of small social organizations or “units”. Inter-unit distances were significantly smaller than those in randomly replaced data, which showed that units associate to form a higher-level social organization or “herd”. Moreover, this herd had a structure where large mixed-sex units were more likely to occupy the center than small mixed-sex units and all-male-units, which were instead on the periphery. These three pieces of evidence regarding the existence of units, unit association, and stable positioning among units strongly indicated a multilevel structure in horse society. The present study contributes to understanding the functions and mechanisms of multilevel societies through comparisons with other social indices and models as well as cross-species comparisons in future studies.

This study only clarified the normal pattern of tablet swallowing under several conditions in healthy subjects, but the results may facilitate comprehensive evaluation and treatment planning in terms of administering medication to dysphagic patients.