Friederike Range

The Evolution of Self-Control

Significance Although scientists have identified surprising cognitive flexibility in animals and potentially unique features of human psychology, we know less about the selective forces that favor cognitive evolution, or the proximate biological mechanisms underlying this process. We tested 36 species in two problem-solving tasks measuring self-control and evaluated the leading hypotheses regarding how and why cognition evolves. Across species, differences in absolute (not relative) brain volume best predicted performance on these tasks. Within primates, dietary breadth also predicted cognitive performance, whereas social group size did not. These results suggest that increases in absolute brain size provided the biological foundation for evolutionary increases in self-control, and implicate species differences in feeding ecology as a potential selective pressure favoring these skills. Cognition presents evolutionary research with one of its greatest challenges. Cognitive evolution has been explained at the proximate level by shifts in absolute and relative brain volume and at the ultimate level by differences in social and dietary complexity. However, no study has integrated the experimental and phylogenetic approach at the scale required to rigorously test these explanations. Instead, previous research has largely relied on various measures of brain size as proxies for cognitive abilities. We experimentally evaluated these major evolutionary explanations by quantitatively comparing the cognitive performance of 567 individuals representing 36 species on two problem-solving tasks measuring self-control. Phylogenetic analysis revealed that absolute brain volume best predicted performance across species and accounted for considerably more variance than brain volume controlling for body mass. This result corroborates recent advances in evolutionary neurobiology and illustrates the cognitive consequences of cortical reorganization through increases in brain volume. Within primates, dietary breadth but not social group size was a strong predictor of species differences in self-control. Our results implicate robust evolutionary relationships between dietary breadth, absolute brain volume, and self-control. These findings provide a significant first step toward quantifying the primate cognitive phenome and explaining the process of cognitive evolution.

The evolution of imitation: what do the capacities of non-human animals tell us about the mechanisms of imitation?

In this paper, we review reports and present new empirical data from studies with marmosets and dogs that address the correspondence problem of imitation research. We focus on the question of how it is possible to transform visual information into matching motor acts. Here, the important issue is not the learning of a complex skill, but determining the copying fidelity of animals at different levels of behavioural organization. As a theoretical framework, we suggest a classification in terms of movement, action and result, which shows a positive relationship between the organizational level of imitation and matching degree. While the monkey studies have provided evidence of very precise copying of movements and, to a lesser degree, of behaviours, the dog studies have provided evidence of action copying and the reproduction of results. In a Do-as-I-do study, a dog attempted to reproduce the results of demonstrated object manipulations at the expense of movement details. Transitive actions were more easily replicated than intransitive ones, and familiarity of actions had a major influence. The discussion of these findings addresses the question of the neuronal mechanisms underlying imitation and whether a single mechanism is sufficient to explain the different levels of copying fidelity.

Dogs’ attention towards humans depends on their relationship, not only on social familiarity

Both in humans and non-human animals, it has been shown that individuals attend more to those they have previously interacted with and/or they are more closely associated with than to unfamiliar individuals. Whether this preference is mediated by mere social familiarity based on exposure or by the specific relationship between the two individuals, however, remains unclear. The domestic dog is an interesting subject in this line of research as it lives in the human environment and regularly interacts with numerous humans, yet it often has a particularly close relationship with its owner. Therefore, we investigated how long dogs (Canis familiaris) would attend to the actions of two familiar humans and one unfamiliar experimenter, while varying whether dogs had a close relationship with only one or both familiar humans. Our data provide evidence that social familiarity by itself cannot account for dogs’ increased attention towards their owners since they only attended more to those familiar humans with whom they also had a close relationship.

The maintenance of traditions in marmosets: individual habit, not social conformity? A field experiment.

Background Social conformity is a cornerstone of human culture because it accelerates and maintains the spread of behaviour within a group. Few empirical studies have investigated the role of social conformity in the maintenance of traditions despite an increasing body of literature on the formation of behavioural patterns in non-human animals. The current report presents a field experiment with free-ranging marmosets (Callithrix jacchus) which investigated whether social conformity is necessary for the maintenance of behavioural patterns within groups or whether individual effects such as habit formation would suffice. Methods Using a two-action apparatus, we established alternative behavioural patterns in six family groups composed of 36 individuals. These groups experienced only one technique during a training phase and were thereafter tested with two techniques available. The monkeys reliably maintained the trained method over a period of three weeks, despite discovering the alternative technique. Three additional groups were given the same number of sessions, but those 21 individuals could freely choose the method to obtain a reward. In these control groups, an overall bias towards one of the two methods was observed, but animals with a different preference did not adjust towards the group norm. Thirteen of the fifteen animals that discovered both techniques remained with the action with which they were initially successful, independent of the group preference and the type of action (Binomial test: exp. proportion: 0.5, p<0.01). Conclusions The results indicate that the maintenance of behavioural patterns within groups 1) could be explained by the first rewarded manipulation and subsequent habit formation and 2) do not require social conformity as a mechanism. After an initial spread of a behaviour throughout a group, this mechanism may lead to a superficial appearance of conformity without the involvement of such a socially and cognitively complex mechanism. This is the first time that such an experiment has been conducted with free-ranging primates.

Individual and sex differences in learning abilities of ravens

Behavioral and physiological characteristics of individuals within the same species have been found to be stable across time and contexts. In this study, we investigated individual differences in learning abilities and object and social manipulation to test for consistency within individuals across different tasks. Individual ravens (Corvus corax) were tested in simple color and position discrimination tasks to establish their learning abilities. We found that males were significantly better in the acquisition of the first discrimination task and the object manipulation task, but not in any of the other tasks. Furthermore, faster learners engaged less often in manipulations of conspecifics and exploration of objects to get access to food. No relationship between object and social manipulation and reversal training were found. Our results suggest that individual differences in regard to the acquisition of new tasks may be related to personalities or at least object manipulation in ravens.

Social attention in keas, dogs, and human children

Understanding animals’ abilities to cooperate with and learn from each other has been an active field of research in recent years. One important basis for all types of social interactions is the disposition of animals to pay attention to each other—a factor often neglected in discussions and experiments. Since attention differs between species as well as between individuals, it is likely to influence the amount and type of information different species and/or observers may extract from conspecifics in any given situation. Here, we carried out a standardized comparative study on attention towards a model demonstrating food-related behavior in keas, dogs and children. In a series of experimental sessions, individuals watched different conspecific models while searching, manipulating and feeding. Visual access to the demonstration was provided by two observation holes, which allowed us to determine exactly how often and for how long observers watched the model. We found profound differences in the factors that influence attention within as well as between the tested species. This study suggests that attention should be incorporated as an important variable when testing species in social situations.

Wolves Are Better Imitators of Conspecifics than Dogs

Domestication is thought to have influenced the cognitive abilities of dogs underlying their communication with humans, but little is known about its effect on their interactions with conspecifics. Since domestication hypotheses offer limited predictions in regard to wolf-wolf compared to dog-dog interactions, we extend the cooperative breeding hypothesis suggesting that the dependency of wolves on close cooperation with conspecifics, including breeding but also territory defense and hunting, has created selection pressures on motivational and cognitive processes enhancing their propensity to pay close attention to conspecifics’ actions. During domestication, dogs’ dependency on conspecifics has been relaxed, leading to reduced motivational and cognitive abilities to interact with conspecifics. Here we show that 6-month-old wolves outperform same aged dogs in a two-action-imitation task following a conspecific demonstration. While the wolves readily opened the apparatus after a demonstration, the dogs failed to solve the problem. This difference could not be explained by differential motivation, better physical insight of wolves, differential developmental pathways of wolves and dogs or a higher dependency of dogs from humans. Our results are best explained by the hypothesis that higher cooperativeness may come together with a higher propensity to pay close attention to detailed actions of others and offer an alternative perspective to domestication by emphasizing the cooperativeness of wolves as a potential source of dog-human cooperation.

Can simple rules account for the pattern of triadic interactions in juvenile and adult female sooty mangabeys

Several experimental and observational studies have investigated the pattern of triadic relationships in primates to demonstrate knowledge of third-party relationships (i.e. an individual recognizes the relationship that exists between two others). To date, all studies have investigated the recognition of third-party relationships in adult animals. Thus, we do not know whether juveniles have knowledge of third-party rank relationships or whether they use a set of simple rules until they learn the rank relationships of all animals within a group. Here, we present the first data on triadic interactions involving juveniles in a rainforest primate, the sooty mangabey, Cercocebus torquatus atys. In a group of 120 individuals, we found that (1) if two groomers were approached by an individual ranking higher than both, the higher-ranking member of the grooming dyad was significantly more likely to stay than the lower-ranking one; (2) when an animal intervened in a conflict, juveniles and adult females were significantly more likely to support the higher-ranking contestant; and (3) juveniles and females solicited help in conflicts significantly more often from potential allies that outranked them and their target than from allies that ranked below them and/or the opponent. Our results suggest that the triadic interactions of juvenile sooty mangabeys resemble triadic interactions reported for adult members of other species of Old World monkeys, but that most of the observed behaviour patterns could be explained by a set of simple rules instead of invoking higher cognitive abilities.

Female but not male dogs respond to a size constancy violation

Differences between sexes in cognitive processes are widespread in humans and permeate many, if not most, cognitive domains. In animal cognition research, however, possible sex differences are still often neglected. Here, we provide striking evidence for a sex-specific response in an object permanence task in domestic dogs (Canis familiaris). Female dogs responded with significantly increased looking times to a violation of expectancy—a ball ‘magically’ changing size while temporarily occluded. By contrast, male dogs, irrespective of their neuter status, did not respond to the size constancy violation. These results indicate that sex differences in basic cognitive processes may extend to mammals in general, and call for increased consideration of possible sex effects when analysing and interpreting data in animal cognition.

Tracking the evolutionary origins of dog-human cooperation: the “Canine Cooperation Hypothesis”

At present, beyond the fact that dogs can be easier socialized with humans than wolves, we know little about the motivational and cognitive effects of domestication. Despite this, it has been suggested that during domestication dogs have become socially more tolerant and attentive than wolves. These two characteristics are crucial for cooperation, and it has been argued that these changes allowed dogs to successfully live and work with humans. However, these domestication hypotheses have been put forward mainly based on dog-wolf differences reported in regard to their interactions with humans. Thus, it is possible that these differences reflect only an improved capability of dogs to accept humans as social partners instead of an increase of their general tolerance, attentiveness and cooperativeness. At the Wolf Science Center, in order to detangle these two explanations, we raise and keep dogs and wolves similarly socializing them with conspecifics and humans and then test them in interactions not just with humans but also conspecifics. When investigating attentiveness toward human and conspecific partners using different paradigms, we found that the wolves were at least as attentive as the dogs to their social partners and their actions. Based on these findings and the social ecology of wolves, we propose the Canine Cooperation Hypothesis suggesting that wolves are characterized with high social attentiveness and tolerance and are highly cooperative. This is in contrast with the implications of most domestication hypotheses about wolves. We argue, however, that these characteristics of wolves likely provided a good basis for the evolution of dog-human cooperation.