Lab cognition going wild: Implementing a new portable touchscreen system in vervet monkeys.

Touchscreen technology has provided researchers with opportunities to conduct well-controlled cognitive tests with captive animals, allowing researchers to isolate individuals, select participants based on specific traits, and control aspects of the environment. In this study, we aimed to investigate the potential utility of touchscreen technology for the study of cognition in wild vervet monkeys. We assessed the viability of touchscreen testing by comparing rates of participation between wild and sanctuary-housed vervets. Additionally, we compared performance on a simple associative learning task in order to verify that wild participants are able to engage meaningfully with a touchscreen task presented in their natural environment. We presented eight groups of vervet monkeys (four wild and four sanctuary groups, totalling 240 individuals) with a portable touchscreen device. The touchscreen displayed tasks in which food rewards could be gained by touching a stimulus displayed on the screen. We assessed individuals' likelihood of interacting with the touchscreen, their frequency of participation, and their performance on a simple associative learning task. We found that sanctuary-housed monkeys were more likely to interact with the touchscreen. Participation in wild vervet monkeys was influenced by sex and age. However, monkeys in the two contexts (sanctuary vs. wild) did not differ in their performance on a simple associative learning task. This study demonstrates that touchscreen technology can be successfully deployed in a population of wild primates. This gives us a starting point to test animal cognition under natural conditions that include varying group composition, environmental challenges and ongoing activities such as foraging, which are challenging to recreate in captivity. While rates of participation were lower than those found in captivity, reasonable sample sizes can be achieved, and wild primates can successfully learn touchscreen tasks in a manner comparable to their captive counterparts.

Socially transmitted diffusion of a novel behavior from subordinate chimpanzees.

Chimpanzees (Pan troglodytes) demonstrate much cultural diversity in the wild, yet a majority of novel behaviors do not become group-wide traditions. Since many such novel behaviors are introduced by low-ranking individuals, a bias toward copying dominant individuals ("rank-bias") has been proposed as an explanation for their limited diffusion. Previous experimental work showed that chimpanzees (Pan troglodytes) preferentially copy dominant over low-rank models. We investigated whether low ranking individuals may nevertheless successfully seed a beneficial behavior as a tradition if there are no "competing" models. In each of four captive groups, either a single high-rank (HR, n = 2) or a low-rank (LR, n = 2) chimpanzee model was trained on one method of opening a two-action puzzle-box, before demonstrating the trained method in a group context. This was followed by 8 hr of group-wide, open-access to the puzzle-box. Successful manipulations and observers of each manipulation were recorded. Barnard's exact tests showed that individuals in the LR groups used the seeded method as their first-choice option at significantly above chance levels, whereas those in the HR groups did not. Furthermore, individuals in the LR condition used the seeded method on their first attempt significantly more often than those in the HR condition. A network-based diffusion analysis (NBDA) revealed that the best supported statistical models were those in which social transmission occurred only in groups with subordinate models. Finally, we report an innovation by a subordinate individual that built cumulatively on existing methods of opening the puzzle-box and was subsequently copied by a dominant observer. These findings illustrate that chimpanzees are motivated to copy rewarding novel behaviors that are demonstrated by subordinate individuals and that, in some cases, social transmission may be constrained by high-rank demonstrators.

"Model age-based" and "copy when uncertain" biases in children's social learning of a novel task.

Theoretical models of social learning predict that individuals can benefit from using strategies that specify when and whom to copy. Here the interaction of two social learning strategies, model age-based biased copying and copy when uncertain, was investigated. Uncertainty was created via a systematic manipulation of demonstration efficacy (completeness) and efficiency (causal relevance of some actions). The participants, 4- to 6-year-old children (N=140), viewed both an adult model and a child model, each of whom used a different tool on a novel task. They did so in a complete condition, a near-complete condition, a partial demonstration condition, or a no-demonstration condition. Half of the demonstrations in each condition incorporated causally irrelevant actions by the models. Social transmission was assessed by first responses but also through children's continued fidelity, the hallmark of social traditions. Results revealed a bias to copy the child model both on first response and in continued interactions. Demonstration efficacy and efficiency did not affect choice of model at first response but did influence solution exploration across trials, with demonstrations containing causally irrelevant actions decreasing exploration of alternative methods. These results imply that uncertain environments can result in canalized social learning from specific classes of model.

The forgotten adaptive social benefits of social learning in animals.

Theoretical and empirical scholars of cultural evolution have traditionally studied social learning strategies, such as conformity, as adaptive strategies to obtain accurate information about the environment, whereas within social psychology there has been a greater focus upon the social consequences of such strategies. Although these two approaches are often used in concert when studying human social learning, we believe the potential social benefits of conformity, and of social learning more broadly, have been overlooked in studies of non-humans. We review evidence from studies of homophily, imitation, and rapid facial mimicry that suggests that behaving like others affords social benefits to non-human animals and that behaviour matching may be deployed strategically to increase affiliation. Furthermore, we review studies of conformity in dispersers, and suggest that forgoing personal information or preferences in favour of those of the new group during immigration may be a strategy to facilitate social integration. We therefore propose that the informational and social functions of conformity apply to humans and animals alike. We use this perspective to generate several interesting research questions to inspire work in this field. For example, under what conditions do animals use informational or social conformity and what role does uncertainty play in social learning in immigrant individuals?

Collective knowledge and the dynamics of culture in chimpanzees.

Social learning in non-human primates has been studied experimentally for over 120 years, yet until the present century this was limited to what one individual learns from a single other. Evidence of group-wide traditions in the wild then highlighted the collective context for social learning, and broader 'diffusion experiments' have since demonstrated transmission at the community level. In the present article, we describe and set in comparative perspective three strands of our recent research that further explore the collective dimensions of culture and cumulative culture in chimpanzees. First, exposing small communities of chimpanzees to contexts incorporating increasingly challenging, but more rewarding tool use opportunities revealed solutions arising through the combination of different individuals' discoveries, spreading to become shared innovations. The second series of experiments yielded evidence of conformist changes from habitual techniques to alternatives displayed by a unanimous majority of others but implicating a form of quorum decision-making. Third, we found that between-group differences in social tolerance were associated with differential success in developing more complex tool use to exploit an increasingly inaccessible resource. We discuss the implications of this array of findings in the wider context of related studies of humans, other primates and non-primate species. This article is part of a discussion meeting issue 'The emergence of collective knowledge and cumulative culture in animals, humans and machines'.

Chimpanzees' behavioral flexibility, social tolerance, and use of tool-composites in a progressively challenging foraging problem.

Behavioral flexibility is a critical ability allowing animals to respond to changes in their environment. Previous studies have found evidence of inflexibility when captive chimpanzees are faced with changing task parameters. We provided two groups of sanctuary-housed chimpanzees with a foraging task in which solutions were restricted over time. Initially, juice could be retrieved from within a tube by hand or by using tool materials, but effective solutions were then restricted by narrowing the tube, necessitating the abandonment of previous solutions and adoption of new ones. Chimpanzees responded flexibly, but one group increased their use of effective techniques to a greater extent than the other. Tool-composite techniques emerged in both groups, but primarily in the more flexible group. The more flexible group also showed higher rates of socio-positive behaviors at the task. In conjunction, these findings support the hypothesis that social tolerance may facilitate the emergence and spread of novel behaviors.

Innovation in chimpanzees.

The study of innovation in non-human animals (henceforth: animals) has recently gained momentum across fields including primatology, animal behaviour and cultural evolution. Examining the rate of innovations, and the cognitive mechanisms driving these innovations across species, can provide insights into the evolution of human culture. Especially relevant to the study of human culture is one of our closest living relatives, the chimpanzee (Pan troglodytes). Both wild and captive chimpanzees demonstrate an impressive ability to innovate solutions to novel problems, but also a striking level of conservatism in some contexts, creating a unique and at times puzzling, picture of animal innovation. Whilst the animal innovation field is rife with potential for expanding our knowledge of human and non-human cognition and problem-solving, it is undermined by a lack of consistency across studies. The field is yet to settle on a definition of the term 'innovation', leading to studies being incomparable across and even within the same species. Here, we fill two gaps in the literature. First, we discuss some of the most prevalent definitions of 'innovation' from different fields, highlighting similarities and differences between them. Secondly, we provide an up-to-date review of accounts of innovations in both wild and captive chimpanzees. We hope this review will provide a resource for researchers interested in the study of innovation in chimpanzees and other animals, as well as emphasising the need for consistency in the way in which innovations are reported.

Chimpanzees (Pan troglodytes) display limited behavioural flexibility when faced with a changing foraging task requiring tool use.

Behavioural flexibility, the ability to alter behaviour in response to environmental feedback, and to relinquish previously successful solutions to problems, is a crucial ability in allowing organisms to adapt to novel environments and environmental change; it is essential to cumulative cultural change. To explore this ability in chimpanzees, 18 individuals (Pan troglodytes) were presented with an artificial foraging task consisting of a tube partially filled with juice that could be reached by hand or retrieved using tool materials to hand. Effective solutions were then restricted in the second phase of the study by narrowing the diameter of the tube, necessitating the abandonment of previously successful solutions. Chimpanzees showed limited behavioural flexibility in comparison to some previous studies, increasing their use of effective techniques, but also continuing to attempt solutions that had been rendered ineffective. This adds to a literature reporting divergent evidence for flexibility (the ability to alter behaviour in response to environmental feedback, and to relinquish previously successful solutions to problems) versus conservatism (a reluctance or inability to explore or adopt novel solutions to problems when a solution is already known) in apes.

A case study of research ethics capacity development in Africa.

Debates about cross-cultural research have sparked growing interest in international research ethics and have initiated collaborative efforts addressing international ethical concerns. This is a case study describing the attempt of the Johns Hopkins Fogarty African Research Ethics Training Program (JHF), recipient of a National Institutes of Health Fogarty International Center (FIC) award, to strengthen research ethics capacity in Africa. The aim of this article is to identify critical elements of an effective research ethics capacity development process through a case study of this particular training program. More specifically, this article describes the JHF program in detail, assesses its outputs during four years, and analyzes its implications of the current model for further growth of this and similar FIC programs. This article assesses the JHF program using materials produced in the first four years of its existence (2001-2004): curriculum materials; application records; informal progress notes and evaluations; transcripts from the trainees' coursework; resumes; and formal progress reports submitted by trainees. The framework used to assess the program is a systems approach, which explores inputs, processes, outputs, and outcomes of the program. The nature and types of inputs changed over time as experience was gained in the program and a continuous improvement in specific processes was implemented. The JHF program has been successful in providing research ethics education and motivating trainees to contribute in the field of research ethics in their countries. How this translates to changes in research ethics in Africa remains to be seen.