Claudio Tennie

Two Key Steps in the Evolution of Human Cooperation

Modern theories of the evolution of human cooperation focus mainly on altruism. In contrast, we propose that humans’ species-unique forms of cooperation—as well as their species-unique forms of cognition, communication, and social life—all derive from mutualistic collaboration (with social selection against cheaters). In a first step, humans became obligate collaborative foragers such that individuals were interdependent with one another and so had a direct interest in the well-being of their partners. In this context, they evolved new skills and motivations for collaboration not possessed by other great apes (joint intentionality), and they helped their potential partners (and avoided cheaters). In a second step, these new collaborative skills and motivations were scaled up to group life in general, as modern humans faced competition from other groups. As part of this new group-mindedness, they created cultural conventions, norms, and institutions (all characterized by collective intentionality), with knowledge of a specific set of these marking individuals as members of a particular cultural group. Human cognition and sociality thus became ever more collaborative and altruistic as human individuals became ever more interdependent.Modern theories of the evolution of human cooperation focus mainly on altruism. In contrast, we propose that humans’ species-unique forms of cooperation—as well as their species-unique forms of cognition, communication, and social life—all derive from mutualistic collaboration (with social selection against cheaters). In a first step, humans became obligate collaborative foragers such that individuals were interdependent with one another and so had a direct interest in the well-being of their partners. In this context, they evolved new skills and motivations for collaboration not possessed by other great apes (joint intentionality), and they helped their potential partners (and avoided cheaters). In a second step, these new collaborative skills and motivations were scaled up to group life in general, as modern humans faced competition from other groups. As part of this new group-mindedness, they created cultural conventions, norms, and institutions (all characterized by collective intentionality), with knowledge of a specific set of these marking individuals as members of a particular cultural group. Human cognition and sociality thus became ever more collaborative and altruistic as human individuals became ever more interdependent.

The nature of prosociality in chimpanzees

An important debate centres around the nature of prosociality in nonhuman primates. Chimpanzees help other individuals in some experimental settings, yet they do not readily share food. One solution to this paradox is that they are motivated to help others provided there are no competing interests. However, benefits to recipients could arise as by-products of testing. Here we report two studies that separate by-product from intended helping in chimpanzees using a GO/NO-GO paradigm. Actors in one group could help a recipient by releasing a food box, but the same action for another group prevented a recipient from being able to get food. We find no evidence for helping—chimpanzees engaged in the test regardless of the effects on their partners. Illusory prosocial behaviour could arise as a by-product of task design.

Conformity cannot be identified based on population-level signatures

Conformist transmission, defined as a disproportionate likelihood to copy the majority, is considered a potent mechanism underlying the emergence and stabilization of cultural diversity. However, ambiguity within and across disciplines remains as to how to identify conformist transmission empirically. In most studies, a population level outcome has been taken as the benchmark to evidence conformist transmission: a sigmoidal relation between individuals’ probability to copy the majority and the proportional majority size. Using an individual-based model, we show that, under ecologically plausible conditions, this sigmoidal relation can also be detected without equipping individuals with a conformist bias. Situations in which individuals copy randomly from a fixed subset of demonstrators in the population, or in which they have a preference for one of the possible variants, yield similar sigmoidal patterns as a conformist bias would. Our findings warrant a revisiting of studies that base their conformist transmission conclusions solely on the sigmoidal curve. More generally, our results indicate that population level outcomes interpreted as conformist transmission could potentially be explained by other individual-level strategies, and that more empirical support is needed to prove the existence of an individual-level conformist bias in human and other animals.

Leaf Surface Roughness Elicits Leaf Swallowing Behavior in Captive Chimpanzees (Pan troglodytes) and Bonobos (P. paniscus), but not in Gorillas (Gorilla gorilla) or Orangutans (Pongo abelii)

Researchers have described apparently self-medicative behaviors for a variety of nonhuman species including birds and primates. Wild chimpanzees, bonobos, and gorillas have been observed to swallow rough leaves without chewing, a behavior proposed to be self-medicative and to aid control of intestinal parasites. Researchers have hypothesized that the presence of hairs on the leaf surface elicits the behavior. We investigated the acquisition and the underlying mechanisms of leaf swallowing. We provided 42 captive great apes (24 chimpanzees, six bonobos, six gorillas, and six orangutans) with both rough-surfaced and hairless plants. None of the subjects had previously been observed to engage in leaf swallowing behavior and were therefore assumed naïve. Two chimpanzees and one bonobo swallowed rough-surfaced leaves spontaneously without chewing them. In a social setup six more chimpanzees acquired the behavior. None of the gorillas or orangutans showed leaf swallowing. Because this behavior occurred in naïve individuals, we conclude that it is part of the behavioral repertoire of chimpanzees and bonobos. Social learning is thus not strictly required for the acquisition of leaf swallowing, but it may still facilitate its expression. The fact that apes always chewed leaves of hairless control plants before swallowing, i.e., normal feeding behavior, indicates that the surface structure of leaves is indeed a determinant for initiating leaf swallowing in apes where it occurs.

Young children spontaneously invent wild great apes’ tool-use behaviours

The variety and complexity of human-made tools are unique in the animal kingdom. Research investigating why human tool use is special has focused on the role of social learning: while non-human great apes acquire tool-use behaviours mostly by individual (re-)inventions, modern humans use imitation and teaching to accumulate innovations over time. However, little is known about tool-use behaviours that humans can invent individually, i.e. without cultural knowledge. We presented 2- to 3.5-year-old children with 12 problem-solving tasks based on tool-use behaviours shown by great apes. Spontaneous tool use was observed in 11 tasks. Additionally, tasks which occurred more frequently in wild great apes were also solved more frequently by human children. Our results demonstrate great similarity in the spontaneous tool-use abilities of human children and great apes, indicating that the physical cognition underlying tool use shows large overlaps across the great ape species. This suggests that humans are neither born with special physical cognition skills, nor that these skills have degraded due to our species’ long reliance of social learning in the tool-use domain.

The island test for cumulative culture in the paleolithic

Early Stone Age artifacts have long been assumed to reflect the material record of communities whose members possessed the ability to transmit ideas, behaviors, and technologies from individual to individual through high-fidelity transmission (i.e., involving teaching and/or imitation), much like humans do today. Recent experimental work has highlighted marked differences between great apes and modern humans in the capacity and/or motivation for some forms of cultural transmission. In particular, high-fidelity mechanisms of social learning, which are thought to underlie the capacity for cumulative culture, appear to be enhanced in – if not unique to – humans. Taken as a group, these experiments suggest it is plausible that a combination of genetic, environmental, and social factors that do not include high-fidelity social learning mediate the “cultures” described for great ape populations to date. It may be that, while the distribution of great ape behavioral variation in time and space is likely affected by low-fidelity social learning (which is widespread in the animal kingdom), the observed variants were invented (i.e., learned) independently by each individual rather than copied from other individuals. Behaviors that do not require high-fidelity transmission between individuals in order to increase in frequency in a population lie within the so-called “zone of latent solutions.” Here, we begin to grapple with the hypothesis that much of the Early Stone Age archaeological record may reflect deeply “canalized” behaviors of hominin toolmakers – those that reside in each individual’s zone of latent solutions – rather than behaviors that necessarily require high-fidelity transmission between individuals. We explore this possibility while eschewing the simplistic notion that variation in stone tool shape, for example, is entirely determined by the genetic variation found in the toolmakers. Instead, we suggest that the variation observed in Early Stone Age artifacts may simply reflect a heavier reliance on behaviors that reside within the zone of latent solutions than on behaviors that make use of high-fidelity social learning. We discuss a thought experiment, called the Island Test, which may be useful for distinguishing hominin behaviors that require high-fidelity transmission from behaviors that do not. We conclude that the Early Stone Age archaeological record is consistent with the possibility that latent solutions explain the behavioral variation inferred from available material culture. Furthermore, we explore reasons why the assumption of high-fidelity transmission associated with Paleolithic industries is difficult to support.

Food washing and placer mining in captive great apes

Sweet potato washing and wheat placer mining in Japanese macaques (Macaca fuscata) are among the most well known examples of local traditions in non-human animals. The functions of these behaviors and the mechanisms of acquisition and spread of these behaviors have been debated frequently. Prompted by animal caretaker reports that great apes [chimpanzees (Pan troglodytes), bonobos (Pan paniscus), gorillas (Gorilla gorilla), and orangutans (Pongo abelii)] at Leipzig Zoo occasionally wash their food, we conducted a study of food washing behaviors that consisted of two parts. In the first part we assessed the current distribution of the behavior on the basis of caretaker reports. In the second (experimental) part, we provided subjects individually with a water basin and two types of food (apples and cereal) that was either clean or covered/mixed with sand. We found that subjects of all species (except gorillas) placed apples in the water before consumption, and that they did so more often when the apples were dirty than when they were clean. Several chimpanzees and orangutans also engaged in behaviors resembling wheat placer mining.

Young children copy cumulative technological design in the absence of action information

The ratchet effect – the accumulation of beneficial changes in cultural products beyond a level that individuals could reach on their own – is a topic of increasing interest. It is currently debated which social learning mechanisms allow for the generation and transmission of cumulative culture. This study focused on transmission, investigating whether 4- to 6-year-old children were able to copy cumulative technological design and whether they could do so without action information (emulation). We adapted the spaghetti tower task, previously used to test for accumulation of culture in human adults. A baseline condition established that the demonstrated tower design was beyond the innovation skills of individual children this age and so represented a culture-dependent product for them. There were 2 demonstration conditions: a full demonstration (actions plus (end-)results) and an endstate- demonstration (end-results only). Children in both demonstration conditions built taller towers than those in the baseline. Crucially, in both demonstration conditions some children also copied the demonstrated tower. We provide the first evidence that young children learn from, and that some of them even copy, cumulative technological design, and that – in line with some adult studies – action information is not always necessary to transmit culture-dependent traits.

The role of redundant information in cultural transmission and cultural stabilization.

Redundant copying has been proposed as a manner to achieve the high-fidelity necessary to pass on and preserve complex traits in human cultural transmission. There are at least 2 ways to define redundant copying. One refers to the possibility of copying repeatedly the same trait over time, and another to the ability to exploit multiple layers of information pointing to the same trait during a single copying event. Using an individual-based model, we explore how redundant copying (defined as in the latter way) helps to achieve successful transmission. The authors show that increasing redundant copying increases the likelihood of accurately transmitting a behavior more than either augmenting the number of copying occasions across time or boosting the general accuracy of social learning. They also investigate how different cost functions, deriving, for example, from the need to invest more energy in cognitive processing, impact the evolution of redundant copying. The authors show that populations converge either to high-fitness/high-costs states (with high redundant copying and complex culturally transmitted behaviors; resembling human culture) or to low-fitness/low-costs states (with low redundant copying and simple transmitted behaviors; resembling social learning forms typical of nonhuman animals). This outcome may help to explain why cumulative culture is rare in the animal kingdom.

Spontaneous reoccurrence of “scooping”, a wild tool-use behaviour, in naïve chimpanzees

Modern human technological culture depends on social learning. A widespread assumption for chimpanzee tool-use cultures is that they, too, are dependent on social learning. However, we provide evidence to suggest that individual learning, rather than social learning, is the driver behind determining the form of these behaviours within and across individuals. Low-fidelity social learning instead merely facilitates the reinnovation of these behaviours, and thus helps homogenise the behaviour across chimpanzees, creating the population-wide patterns observed in the wild (what here we call “socially mediated serial reinnovations”). This is the main prediction of the Zone of Latent Solutions (ZLS) hypothesis. This study directly tested the ZLS hypothesis on algae scooping, a wild chimpanzee tool-use behaviour. We provided naïve chimpanzees (n = 14, Mage = 31.33, SD = 10.09) with ecologically relevant materials of the wild behaviour but, crucially, without revealing any information on the behavioural form required to accomplish this task. This study found that naïve chimpanzees expressed the same behavioural form as their wild counterparts, suggesting that, as the ZLS theory predicts, individual learning is the driver behind the frequency of this behavioural form. As more behaviours are being found to be within chimpanzee’s ZLS, this hypothesis now provides a parsimonious explanation for chimpanzee tool cultures.