Experimental approaches to the study of culture in primates

First paragraph: The topic of culture in nonhumans has received a lot of attention in recent years, and perhaps this is not surprising. The idea that other animals - apes, monkeys, and maybe even rats and fish - may show a semblance of a trait that we consider to be so fundamentally human, is both exciting and controversial. Dialogue between enthusiastic advocates of animal culture on the one hand, and wary sceptics on the other, has therefore generated a fruitful intellectual atmosphere, and spawned a broad, rich, scientific literature on the topic

Cultural Evolution and Perpetuation of Arbitrary Communicative Conventions in Experimental Microsocieties

Previous studies have shown that iconic graphical signs can evolve into symbols through repeated usage within dyads and interacting communities. Here we investigate the evolution of graphical signs over chains of participants. In these chains (or "replacement microsocieties"), membership of an interacting group changed repeatedly such that the most experienced members were continually replaced by naive participants. Signs rapidly became symbolic, such that they were mutually incomprehensible across experienced members of different chains, and new entrants needed to learn conventionalised meanings. An objective measure of graphical complexity (perimetric complexity) showed that the signs used within the microsocieties were becoming progressively simplified over successive usage. This is the first study to show that the signs that evolve in graphical communication experiments can be transmitted to, and spontaneously adopted by, naive participants. This provides critical support for the view that human communicative symbols could have evolved culturally from iconic representations

Human Cumulative Culture in the Laboratory: Effects of (Micro) Population Size

Traditionally, experiments on social learning (both in humans and nonhumans) involve dyads, with an experimenter or experimenter-trained conspecific serving as the demonstrator and the participant as the observer. But social learning in nature often involves multiple potential models, and the models themselves were once learners. We discuss our studies of social learning in adult humans in interactive group settings in the absence of formal demonstrations by experimenters, which track transmission over multiple learner generations. In these experiments we find evidence for cumulative learning over generations. This has allowed us to manipulate learning conditions in order to test hypotheses regarding the necessary conditions for cumulative culture. We also report results from a further experiment using similar methods, which compares conditions of varying cohort size. Participants were given the task to build a paper airplane to fly as far as possible. Contrary to expectations, there was no advantage for larger cohort sizes, in terms of the cumulative effects observed

Artificially generated cultural variation between two groups of captive colobus monkeys, Colobus guereza kikuyuensis

The majority of studies of social learning in primates have tested subjects in isolation and investigated the effects of learning over very short periods of time. We aimed to test for social learning in two social groups of colobus monkeys, Colobus guereza kikuyuensis. Subjects were shown video footage of familiar monkeys either pushing or pulling a plastic flap to obtain a food reward, while they were given simultaneous access to the same apparatus. Action frequencies showed a significant difference between the two groups, with the pull group performing a higher proportion of pulls to pushes, compared with the push group. Copying persisted even in later sessions during which the demonstration footage was not being shown. We conclude that we successfully generated two contrasting behavioural traditions in these groups of monkeys. We do not know how long this contrast in behaviour would have persisted had we been able to continue testing for an even longer period of time, but further studies using similar designs and even longer test periods would have the power to confirm whether stable behavioural variation can be sustained between groups of monkeys, supported by social transmission

Human teaching and cumulative cultural evolution

Although evidence of teaching behaviour has been identified in some nonhuman species, human teaching appears to be unique in terms of both the breadth of contexts within which it is observed, and in its responsiveness to needs of the learner. Similarly, cultural evolution is observable in other species, but human cultural evolution appears strikingly distinct. This has led to speculation that the evolutionary origins of these capacities may be causally linked. Here we provide an overview of contrasting perspectives on the relationship between teaching and cultural evolution in humans, and briefly review previous research which suggests that cumulative culture (here meaning cultural evolution featuring a trend towards improving functionality) can occur without teaching. We then report the results of a novel experimental study in which we investigated how the benefits of teaching may depend on the complexity of the skill to be acquired. Participants were asked to tie knots of varying complexity. In our Teaching condition, opportunities to interact with an experienced partner aided transmission of the most complex knots, but not simpler equivalents, relative to exposure to completed products alone (End State Only condition), and also relative to information about the process of completion (Intermediate States condition). We conclude by considering the plausibility of various accounts of the evolutionary relationship between teaching and cultural evolution in humans

Experimental approaches to studying cumulative cultural evolution

In humans, cultural traditions often change in ways which increase efficiency and functionality. This process, widely referred to as cumulative cultural evolution, sees beneficial traits preferentially retained, and it is so pervasive that we may be inclined to take it for granted. However, directional change of this kind appears to distinguish human cultural traditions from behavioural traditions that have been documented in other animals. Cumulative culture is therefore attracting an increasing amount of attention within psychology, and researchers have begun to develop methods of studying this phenomenon under controlled conditions. These studies have now addressed a number of different questions, including which learning mechanisms may be implicated, and how the resulting behaviours may be influenced by factors such as population structure. The current article provides a synopsis of some of these studies, and highlights some of the unresolved issues in this field

Cultural diffusion in humans and other animals

Recent years have seen an enormous expansion and progress in studies of the cultural diffusion processes through which behaviour patterns, ideas and artifacts are transmitted within and between generations of humans and other animals. The first of two main approaches focuses on identifying, tracing and understanding cultural diffusion as it naturally occurs, an essential foundation to any science of culture. This endeavor has been enriched in recent years by sophisticated statistical methods and surprising new discoveries particularly in humans, other primates and cetaceans. This work has been complemented by a growing corpus of powerful, purpose-designed cultural diffusion experiments with captive and natural populations that have facilitated the rigorous identification and analysis of cultural diffusion in species from insects to humans

Identifying innovation in laboratory studies of cultural evolution: rates of retention and measures of adaptation

In recent years, laboratory studies of cultural evolution have become increasingly prevalent as a means of identifying and understanding the effects of cultural transmission on the form and functionality of transmitted material. The data sets generated by these studies may provide insights into the conditions encouraging, or inhibiting, high rates of innovation, as well as the effect that this has on measures of adaptive cultural change. Here we review recent experimental studies of cultural evolution with a view to elucidating the role of innovation in generating observed trends. We first consider how tasks are presented to participants, and how the corresponding conceptualisation of task success is likely to influence the degree of intent underlying any deviations from perfect reproduction. We then consider the measures of interest used by the researchers to track the changes that occur as a result of transmission, and how these are likely to be affected by differing rates of retention. We conclude that considering studies of cultural evolution from the perspective of innovation provides valuable insights which help to clarify important differences in research designs, which have implications for the likely effects of variation in retention rates on measures of cultural adaptatio

Opportunity to assimilate and pressure to discriminate can generate cultural divergence in the laboratory

Formal models of cultural evolution have illustrated circumstances under which behavioral traits that have no inherent advantage over others can undergo positive selection pressure. One situation in which this may occur is when the behavior functions as a social marker, and there is pressure to identify oneself as a member of a particular group. Our aim in the current study was to determine whether participants organized into subpopulations could effectively exploit variation in a completely novel behavior to advertise themselves as belonging to a particular subpopulation, such that discrimination between in-group and out-group members was possible and subpopulations exhibited increasing distinctiveness. Eighty participants took part, organized into four subpopulations, each comprised of five four-member generations. They each completed a tower-building task, used in previous experimental studies of cultural evolution. An incentive payment structure was imposed with the aim of motivating participants to advertise themselves as belonging to a particular subpopulation, and to distinguish in-group members from members of other subpopulations. The first generation were exposed to photographs of randomly-assigned "seed" towers, and later generations were exposed to photographs of the towers built by the members of the previous generation of their own subpopulation. Participants were able to discriminate towers built by in-group members of the same generation, from towers built by out-group members. Over generations, tower designs evolved such that they were increasingly identifiable as belonging to a particular subpopulation. Arbitrary traits, which had no prior advantage, became associated with group membership, providing empirical support for theoretical models

End State Copying by Humans (Homo sapiens): Implications for a Comparative Perspective on Cumulative Culture

It has been proposed that the uniqueness of human cumulative culture may be attributable to humans' greater orientation towards copying the process of behavior (imitation), compared with the products (emulation), resulting in particularly high fidelity transmission. Following on from previous work indicating that adult human participants can exhibit cumulative learning on the basis of product copying alone, we now investigate whether such learning involves high fidelity transmission. Eighty adult human participants were presented with a task previously shown to elicit cumulative learning under experimental conditions, which involved building a tower from spaghetti and modeling clay. Each participant was shown two completed towers, ostensibly built by previous participants, but actually built to pre-specified designs by the experimenter. This end state information was provided either in the form of photographs, or the presence of actual towers. High fidelity matching to these end states was apparent in both demonstration conditions, even for a design that was demonstrably suboptimal with regard to the goal of the task (maximizing tower height). We conclude that, although high fidelity transmission is likely to be implicated in cumulative culture, action copying is not always necessary for this to occur. Furthermore, since chimpanzees apparently copy behavioral processes and well as products, and also transmit behavior with high fidelity, the stark absence of unequivocal examples of cumulative culture in nonhumans may be attributable to factors other than imitative ability.Research funded by ESR