On experimental tests for studying altruism in capuchin monkeys.

Altruism is often considered as the ultimate form of prosociality and is defined as any act that benefits others without direct benefit to the actor. Many nonhuman species have been reported to express different forms of altruism, although their expression in experimental studies is highly dependent on the paradigm used. Tufted capuchin monkeys are one of the most studied species; however, the evidence for altruism in this species remains inconclusive. This study aimed to investigate whether two paradigms, adapted from those in which great apes have shown altruism, could be useful for revealing signs of altruistic capabilities in capuchins. Pairs of monkeys were tested in two experiments involving a similar mechanism but with different costs to acting altruistically. The first used a more costly operant sharing task in which an operator could unlock a door to allow a recipient to enter the room and share his food. The second consisted of a less costly helping task, in which the operator's food was secured but he could help the recipient to get other food that was in a locked container. The results suggest that capuchins, although apparently unwilling to share their food in a costly operant situation, might altruistically help selected recipients, in response to requesting by the latter. While our small sample size along with procedural limitations preclude firm conclusions, we discuss how further ameliorations of our tasks could further contribute to the study of altruistic capacities in primates.

Processing of self-initiated sound motion in the human brain.

Interacting with objects in our environment usually leads to audible noise. Brain responses to such self-initiated sounds have been shown to be attenuated, in particular the so-called N1 component measured with electroencephalography (EEG). This attenuation has been proposed to be the effect of an internal forward model that allows for cancellation of the sensory consequences of a motor command. In the current study we asked whether the attenuation due to self-initiation of a sound also affects a later event-related potential - the so-called motion-onset response - that arises in response to moving sounds. To this end, volunteers were instructed to move their index fingers either left or rightward which resulted in virtual movement of a sound either to the left or to the right. In Experiment 1, sound motion was induced with in-ear head-phones by shifting interaural time and intensity differences and thus shifting the intracranial sound image. We compared the motion-onset responses under two conditions: a) congruent, and b) incongruent. In the congruent condition, the sound image moved in the direction of the finger movement, while in the incongruent condition sound motion was in the opposite direction of the finger movement. Clear motion-onset responses with a negative cN1 component peaking at about 160 ms and a positive cP2 component peaking at about 230 ms after motion-onset were obtained for both the congruent and incongruent conditions. However, the motion-onset responses did not significantly differ between congruent and incongruent conditions in amplitude or latency. In Experiment 2, in which sounds were presented with loudspeakers, we observed attenuation for self-induced versus externally triggered sound motion-onset, but again, there was no difference between congruent and incongruent conditions. In sum, these two experiments suggest that the motion-onset response measured by EEG can be attenuated for self-generated sounds. However, our result did not indicate that this attenuation depended on congruency of action and sound motion direction.

Investigating reactions of squirrel monkeys (Saimiri sciureus) towards unequal food distributions in a tray-pulling paradigm.

The fact that squirrel monkeys do not routinely cooperate in the wild has been proposed to explain their failure to show disadvantageous inequity aversion (i.e., negative reactions when receiving less than a partner) in an experimental exchange. Here we assessed whether the use of a tray-pulling paradigm, allowing for a larger variety of unequal testing situations, would bring additional insights into inequity aversion in this species. Squirrel monkeys were tested in pairs in which only the donor could pull a tray baited with food to within reach of itself and a recipient. Using pairs with different social relationships, we examined donors' frequencies of pulling both in the presence and absence of a recipient, as well as across three different food distributions: equal, qualitative inequity (higher-value reward for the recipient), and quantitative inequity (no food reward for the donor). Results showed that female donors pulled the tray less often in the quantitative inequity condition with an out-group female recipient than when alone. However, such discrimination was not observed when females were with female in-group and male out-group recipients. By contrast, male donors did not adjust their pull frequencies according to the recipient's presence or identity (female and male out-group recipients). These results point towards possible disadvantageous inequity aversion in female squirrel monkeys. However, alternative hypotheses such as increased arousal caused by out-group female recipients cannot be ruled out. We discuss the data in line with major theories of inequity aversion and cooperation in primates.

Trading of dynamic interaural time and level difference cues and its effect on the auditory motion-onset response measured with electroencephalography.

Interaural time (ITD) and level differences (ILD) constitute the two main cues for sound localization in the horizontal plane. Despite extensive research in animal models and humans, the mechanism of how these two cues are integrated into a unified percept is still far from clear. In this study, our aim was to test with human electroencephalography (EEG) whether integration of dynamic ITD and ILD cues is reflected in the so-called motion-onset response (MOR), an evoked potential elicited by moving sound sources. To this end, ITD and ILD trajectories were determined individually by cue trading psychophysics. We then measured EEG while subjects were presented with either static click-trains or click-trains that contained a dynamic portion at the end. The dynamic part was created by combining ITD with ILD either congruently to elicit the percept of a right/leftward moving sound, or incongruently to elicit the percept of a static sound. In two experiments that differed in the method to derive individual dynamic cue trading stimuli, we observed an MOR with at least a change-N1 (cN1) component for both the congruent and incongruent conditions at about 160-190 ms after motion-onset. A significant change-P2 (cP2) component for both the congruent and incongruent ITD/ILD combination was found only in the second experiment peaking at about 250 ms after motion onset. In sum, this study shows that a sound which - by a combination of counter-balanced ITD and ILD cues - induces a static percept can still elicit a motion-onset response, indicative of independent ITD and ILD processing at the level of the MOR - a component that has been proposed to be, at least partly, generated in non-primary auditory cortex.

Third-party social evaluations of humans by monkeys and dogs.

Developmental psychologists are increasingly interested in young children's evaluations of individuals based on third-party interactions. Studies have shown that infants react negatively to agents who display harmful intentions toward others, and to those who behave unfairly. We describe experimental studies of capuchin monkeys' and pet dogs' differential reactions to people who are helpful or unhelpful in third-party contexts, and monkeys' responses to people who behave unfairly in exchanges of objects with a third party. We also present evidence that capuchin monkeys monitor the context of failures to help and violations of reciprocity, and that intentionality is one factor underlying their social evaluations of individuals whom they see interacting with others. We conclude by proposing some questions for studies of nonhuman species' third party-based social evaluations.

Microsatellite Polymorphisms Adjacent to the Oxytocin Receptor Gene in Domestic Cats: Association with Personality?

A growing number of studies have explored the oxytocin system in humans and non-human animals, and some have found important genetic polymorphisms in the oxytocin receptor gene (OXTR) associated with the bonding system, social behaviors, and personality in several species. Although single nucleotide polymorphisms in OXTR have been well-examined in various species, microsatellites (or short tandem repeats) adjacent to OXTR have rarely been studied, despite some suggestions that microsatellite polymorphisms near genes might play a role in genetic transcription and translation. In this study, we surveyed microsatellites in the upstream, intron, and downstream regions of OXTR in domestic cats (Felis catus). We succeeded in amplifying 5 out of 10 regions, and recognized these five regions as polymorphic. We compared allele frequencies in these five regions between mongrel cats in Japan (n = 100) and cats of 10 pure breeds (n = 40). There were significant differences in allele frequencies between the two populations in all microsatellite regions. Additionally, the owners of mongrel cats answered a comprehensive personality questionnaire, and factor analysis extracted four factors (Openness, Friendliness, Roughness, and Neuroticism). We examined the association between the microsatellite genotypes, age, sex, neutering status, and personality scores. Compared to their counterparts, younger cats tended to score higher on Openness, male cats scored higher on Friendliness, and female and neutered cats scored higher on Roughness. When we divided the sample into three groups depending on the length of alleles, we found a marginally significant association between Friendliness and MS3. Additionally, we found a sex-mediated effect of genotypes in MS4 on Friendliness, resulting in different effects on females and males. Our findings that mongrel cats had longer alleles in MS3 and MS4 than purebred cats, and that those cats tended to score higher on Friendliness, supported the previous findings. However, future studies such as comparison between purebred cats with apparently different origin or personality are required to determine the association of genetic variants in the OXTR with personality.

Evaluation of third-party reciprocity by squirrel monkeys (Saimiri sciureus) and the question of mechanisms.

Social evaluation during third-party interactions emerges early in human ontogeny, and it has been shown in adult capuchin monkeys who witness violations of reciprocity in object exchanges: Monkeys were less inclined to accept food from humans who refused to reciprocate with another human. A recent study reporting similar evidence in marmoset monkeys raised the possibility that such evaluations might be based on species' inherent cooperativeness. We tested a species not renowned for cooperativeness-squirrel monkeys-using the procedure used with marmosets and found a similar result. This finding rules out any crucial role for cooperative tendencies in monkeys' responses to unfair exchanges. We then tested squirrel monkeys using procedures more similar to those used in the original study with capuchins. Squirrel monkeys again accepted food less frequently from non-reciprocators, but unlike capuchins, they also strongly preferred reciprocators. We conclude that neither squirrel monkeys nor marmoset monkeys engaged in emotional bookkeeping of the type that probably underlies social evaluation in capuchin monkeys; instead, they employed one or more simple behavioral rules. Further comparative studies are required to clarify the mechanisms underlying social evaluation processes across species.