Social influences on ant-dipping acquisition in the wild chimpanzees (Pan troglodytes verus) of Bossou,Guinea, West Africa

We currently have little understanding of the influence of learning opportunity, whether social or environmental, and maternal role on tool-use acquisition in young wild chimpanzees. This study aims to fill this gap by focusing on the acquisition of ant-dipping among chimpanzees of Bossou, Guinea. Ant-dipping is a hazardous tool-use behaviour aimed at army ants (Dorylus spp.). Bossou chimpanzees target these ants both at nests (high risk) and trails (low risk) and employ two techniques to consume them: direct mouthing and pull-through. We present data for 13 mother–offspring pairs (1–10 years old). Mothers with young ≤5 years old dipped significantly more often at trails than at nests, thus minimizing the risk posed to themselves and their young. Infants thus benefited from better conditions to observe and practice ant-dipping. Mothers also varied greatly in their percent time spent ant-dipping and offspring differed in their learning opportunity. Our results suggest that high opportunity young started to observe and perform ant-dipping sooner and were better at ant-dipping than low opportunity young. Although mothers and weaned offspring correlated positively in their percent time spent dipping and proficiency, they did not match in technique used or tool length. Finally, we propose that the learning trajectory of young may predict individual and sex differences in adulthood. This study demonstrates the important role of mothers and learning opportunity in the acquisition of a hazardous tool-use behaviour and suggests that chimpanzee material culture is a product of a complex interaction between social processes and ecological factors.     

Acquisition and memory of sequence order in young and adult chimpanzees (Pan troglodytes)

There is no research about age difference in the process of sequential learning in non-human primates. Is there any difference between young and adults in sequential learning process? Six chimpanzees (Pan troglodytes), 3 young and 3 adults, learned the Arabic numeral sequence 1 to 9 by touching the numerals on a touch-screen monitor in ascending order. Initially, the sequence always started with the numeral 1, i.e. ‘start-fixed task’. Training began with the sequence 1–2, 1–2–3, and continued sequentially up to 1–2–3–4–5–6–7–8–9. Later, the subjects were introduced to sequences that started with a random numeral, but always ended with 9, i.e. ‘end-fixed task’. Performance in the end-fixed task was worse relative to the familiar start-fixed task. After training with various sequences of adjacent numerals, the subjects were given a transfer test for the non-adjacent numerals. The results suggested that all chimpanzees indeed mastered sequential ordering, and although there was no fundamental difference in the acquisition process between the two age groups, there was a significant age difference in memory capacity. Based on their knowledge of sequential ordering, the subjects were then asked to perform a masking task in which once a subject touched the lowest numeral, the other numeral(s) turned to white squares. Performance of the masking task by young chimpanzees was better than that of adults in accuracy and degree of difficulty (number of numerals). Taken together, these data clearly demonstrate a similarity among subjects in the way chimpanzees acquire knowledge of sequential order regardless of age differences in sequential learning. Moreover, they reveal that once knowledge of sequential order is established, it can be a good index used to evaluate memory capacity in young and adult chimpanzees.

     

Personality predicts innovation and social learning in children: Implications for cultural evolution

Innovation and social learning are the pillars of cultural evolution, allowing cultural behaviours to cumulatively advance over generations. Yet, little is known about individual differences in the use of social and asocial information. We examined whether personality influenced 7-11-year-old children's (N = 282) propensity to elect to observe others first or independently generate solutions to novel problems. Conscientiousness was associated with electing for no demonstrations, while agreeableness was associated with opting for demonstrations. For children receiving demonstrations, openness to experience consistently predicted deviation from observed methods. Children who opted for no demonstrations were also more likely than those opting for demonstrations to exhibit tool manufacture on an innovation challenge and displayed higher creativity, as measured by an alternate uses task. These results highlight how new cultural traditions emerge, establish and advance by identifying which individuals generate new cultural variants in populations and which are influential in the diffusion of these variants, and help reduce the apparent tension within the ‘ratchet’ of cumulative culture.     

The evolution of prestige: Perspectives and hypotheses from comparative studies

In this review, we reorganize the concept and highlight the importance of prestige in humans and non-human animals by introducing key characteristics of dominance and prestige and related theories. Previous studies with non-human animals have mainly focused on dominance, presuming prestige as a human-unique social trait. However, to deepen our understanding of the evolution of prestige, comparative studies with non-human animals, especially our evolutionary closest relatives, chimpanzees and bonobos, are essential. We propose the direction of future studies to investigate how prestige has emerged as a viable strategy for gaining social rank while diverging from dominance, which will establish a foundation for investigating the impact of prestige on propensities towards large-scale cooperation and cumulative culture which are still considered unique to humans. Such comparative viewpoints on prestige, along with some hypotheses of species differences, will provide powerful guidance for understanding the evolution of social hierarchies.     

Even simple forms of social learning rely on intention attribution in marmoset monkeys (Callithrix jacchus)

Intention attribution guides the cognitively most demanding forms of social learning, such as imitation, thereby scaffolding cumulative cultural evolution. However, it is not thought to be necessary for more basic forms of social learning. Here we present evidence that in marmoset monkeys (Callithrix jacchus) even most basic forms of social learning such as enhancement depend on intention attribution. Marmosets perceived the behavior of a conspecific and a conspecific-like robot, but not that of a moving black box, as goal directed. Their subsequent choice behavior was shaped by social facilitation and stimulus enhancement, that is, by very simple forms of social learning, but only when exposed to the conspecific and robot, which they previously had perceived as intentional agents. We discuss the implications of this finding for contemporary debates about social learning, including emulation learning and ghost control studies, the necessity of goal-directed copying for cumulative cultural evolution, and the limits of current classification systems of social learning for the evolution of social and asocial learning.     

Conservatism and “copy-if-better” in chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>)

Social learning is predicted to evolve in socially living animals provided the learning process is not random but biased by certain socio-ecological factors. One bias of particular interest for the emergence of (cumulative) culture is the tendency to forgo personal behaviour in favour of relatively better variants observed in others, also known as the “copy-if-better” strategy. We investigated whether chimpanzees employ copy-if-better in a simple token-exchange paradigm controlling for individual and random social learning. After being trained on one token-type, subjects were confronted with a conspecific demonstrator who either received the same food reward as the subject (control condition) or a higher value food reward than the subject (test condition) for exchanging another token-type. In general, the chimpanzees persisted in exchanging the token-type they were trained on individually, indicating a form of conservatism consistent with previous studies. However, the chimpanzees were more inclined to copy the demonstrator in the test compared to the control condition, indicating a tendency to employ a copy-if-better strategy. We discuss the validity of our results by considering alternative explanations and relate our findings to the emergence of cumulative culture.     

Social learning in pig-tailed macaques (Macaca nemestrina) and adult humans (Homo sapiens) on a two-action artificial fruit

An artificial fruit (AF) was used to test for social learning in pig-tailed macaques (Macaca nemestrina) and adult humans (Homo sapiens). A monkey demonstrator opened the AF, showing alternative methods to 2 groups of cage mates. Video films of the monkey demonstrations were presented to adult humans. Compared with chimpanzees and children, the macaques watched the demonstrations significantly less and in a much more sporadic manner. They also produced only very weak and transitory evidence of social learning. In contrast, the adult humans performed as one might expect of optimum imitators, even producing evidence of components of a "ratchet effect."     

Slow development of vocal sequences through ontogeny in wild chimpanzees (Pan troglodytes verus)

The development of the unique, hierarchical, and endless combinatorial capacity in a human language requires neural maturation and learning through childhood. Compared with most non-human primates, where combinatorial capacity seems limited, chimpanzees present a complex vocal system comprising hundreds of vocal sequences. We investigated how such a complex vocal system develops and the processes involved. We recorded 10,929 vocal utterances of 98 wild chimpanzees aged 0–55 years, from Taï National Park, Ivory Coast. We developed customized Generalized non-Linear Models to estimate the ontogenetic trajectory of four structural components of vocal complexity: utterance length, diversity, probability of panting (requiring phonation across inhalation and exhalation), and probability of producing two adjacent panted units. We found chimpanzees need 10 years to reach adult levels of vocal complexity. In three variables, the steepest increase coincided with the age of first non-kin social interactions (2–5 years), and plateaued in sub-adults (8–10 years), as individuals integrate into adult social life. Producing two adjacent panted units may require more neuromuscular coordination of the articulators, as its emergence and steepest increase appear later in development. These results suggest prolonged maturational processes beyond those hitherto thought likely in species that do not learn their vocal repertoire. Our results suggest that multifaceted ontogenetic processes drive increases in vocal structural complexity in chimpanzees, particularly increases in social complexity and neuro-muscular maturation. As humans live in a complex social world, empirical support for the “social complexity hypothesis” may have relevance for theories of language evolution.

Research Highlights

• Chimpanzees need around 10 years to develop the vocal structural complexity present in the adult repertoire, way beyond the age of emergence of every single vocal unit.
• Multifaceted ontogenetic processes may drive increases in vocal structural complexity in chimpanzees, particularly increases in social complexity and neuro-muscular maturation.
• Non-linear increases in vocal complexity coincide with social developmental milestones.
• Vocal sequences requiring rapid articulatory change emerge later than other vocal sequences, suggesting neuro-muscular maturational processes continue through the juvenile years.

     

Neonatal imitation in chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>) tested with two paradigms

Primate species differ in their imitative performance, perhaps reflecting differences in imitative capacity. The developmentally earliest form of imitation in humans, neonatal imitation, occurs in early interactions with social partners, and may be a more accurate index of innate capacity than imitation of actions on objects, which requires more cognitive ability. This study assessed imitative capacity in five neonatal chimpanzees, within a narrow age range (7–15 days of age), by testing responses to facial and vocal actions with two different test paradigms (structured and communicative). Imitation of mouth opening was found in both paradigms. In the communicative paradigm, significant agreement was found between infant actions and demonstrations. Additionally, chimpanzees matched the sequence of three actions of the TC model, but only on the second demonstration. Newborn chimpanzees matched more modeled actions in the communicative test than in the structured paradigm. These performances of chimpanzees, at birth, are in agreement with the literature, supporting a conclusion that imitative capacity is not unique to the human species. Developmental histories must be more fully considered in the cross-species study of imitation, as there is a greater degree of innate imitative capacity than previously known. Socialization practices interact with innate and developing competencies to determine the outcome of imitation tests later in life.     

Copying results and copying actions in the process of social learning: chimpanzees (Pan troglodytes) and human children (Homo sapiens)

There is currently much debate about the nature of social learning in chimpanzees. The main question is whether they can copy others' actions, as opposed to reproducing the environmental effects of these actions using their own preexisting behavioral strategies. In the current study, chimpanzees (Pan troglodytes) and human children (Homo sapiens) were shown different demonstrations of how to open a tube-in both cases by a conspecific. In different experimental conditions, demonstrations consisted of (1) action only (the actions necessary to open the tube without actually opening it); (2) end state only (the open tube, without showing any actions); (3) both of these components (in a full demonstration); or (4) neither of these components (in a baseline condition). In the first three conditions subjects saw one of two different ways that the tube could open (break in middle; caps off ends). Subjects' behavior in each condition was assessed for how often they opened the tube, how often they opened it in the same location as the demonstrator, and how often they copied the demonstrator's actions or style of opening the tube. Whereas chimpanzees reproduced mainly the environmental results of the demonstrations (emulation), human children often reproduced the demonstrator's actions (imitation). Because the procedure used was similar in many ways to the procedure that Meltzoff (Dev Psych 31:1, 1995) used to study the understanding of others' unfulfilled intentions, the implications of these findings with regard to chimpanzees' understanding of others' intentions are also discussed.     

Social learning of nut-cracking behavior in East African sanctuary-living chimpanzees (Pan troglodytes schweinfurthii)

Nut cracking is restricted to communities of wild chimpanzees living in West Africa, suggesting it is an example of a socially transmitted tradition. Detailed study of the acquisition of nut cracking in wild chimpanzees is consistent with this conclusion. However, only 2, small-scale experiments have been carried out in captivity to explore the role of social transmission in the acquisition of this behavior. The study presented here does this with a comparatively larger, statistically viable sample of 11 sanctuary-living chimpanzees (Pan troglodytes schweinfurthii), permitting both between- and within-subjects experimental manipulations. Results confirmed that nut cracking can be acquired in a matter of days by social learning, but only in chimpanzees 3 to 4 years old and older. Direct comparisons are made with a study carried out in the wild, revealing striking similarities in developmental profiles.     

AGE DIFFERENCES IN CUMULATIVE LEARNING OF MEANINGFUL AND MEANINGLESS MATERIAL

Age differences in cumulative learning were explored using a technique developed by Hebb. In the first experiment Finnish subjects recalled messages of English digits and in two subsequent experiments English subjects recalled messages of Finnish and of English digits. The finding from the first experiment was that young subjects exhibited cumulative learning while old subjects did not but it was concluded on the basis of the two further experiments that both age groups could show cumulative learning under the conditions used. An age difference in immediate memory for meaningful and meaningless material is noted and the possibility of increasing vulnerability with age to different forms of interference is discussed.     

Leaf swallowing behavior in chimpanzees (Pan troglodytes): biased learning and the emergence of group level cultural differences

Demonstrating the ability to ‘copy’ the behavior of others is an important aspect in determining whether social learning occurs and whether group level differences in a given behavior represent cultural differences or not. Understanding the occurrence of this process in its natural context is essential, but can be a daunting task in the wild. In order to test the social learning hypothesis for the acquisition of leaf swallowing (LS), a self-medicative behavior associated with the expulsion of parasites, we conducted semi-naturalistic experiments on two captive groups of parasite-free, naïve chimpanzees (Pan troglodytes). Individuals in the group were systematically provided appropriate stimuli (rough hispid leaves) identical to those used by chimpanzees in the wild. Individuals initially responded in a variety of ways, ranging from total aversion to normal chewing and swallowing. Over time, however, the two groups adopted different variants for inserting and folding the leaves in the mouth prior to swallowing them (complete and partial LS), following the specific method spontaneously displayed by the first and primary LS models in their respective groups. These variants were similar to LS displayed by chimpanzees in the wild. Using the option-bias method, we found evidence for social learning leading to group-level biased transmission and group-level stabilization of these two variants. This is the first report on two distinct cultural variants innovated in response to the introduction of natural stimuli that emerged and spread spontaneously and concurrently within two adjacent groups of socially housed primates. These observations support the assertion that LS may reflect a generalized propensity for ingesting rough hispid leaves, which can be socially induced and transmitted within a group. Ingesting an adequate number of these leaves induces increased gut motility, which is responsible for the subsequent expulsion of particular parasite species in the wild. Cultural transmission and maintenance of LS within a group and associative learning by the individual of the positive consequences of this otherwise non-nutritive mode of ingestion is proposed to be the pivotal link between this feeding propensity and its maintenance as a self-medicative behavior by great apes in the wild.     

The interaction of observational learning with overt practice: effects on motor skill learning

This study explored various methods of combining observational learning via demonstration with the effects of overt practice for learning a discrete action pattern. Three groups were compared that varied by the timing of demonstration in relation to practice. An all-pre-practice demonstration group viewed 10 pre-practice videotape demonstrations of an expert performing the skill, and then engaged in practice. An interspersed demonstration group viewed one pre-practice demonstration, then initiated practice on the skill. Every three attempts, practice was halted while participants viewed another demonstration, with this pattern repeated throughout acquisition. A combination demonstration group experienced elements of each schedule by viewing five demonstrations prior to practice, then five more once practice had begun (one every three attempts) so that modeling was completed by mid-acquisition. Ratings of form and accuracy were assessed in an acquisition phase, an immediate retention test, and a 48-h retention test. Group main effects for form scores were detected in acquisition, immediate, and 48-h retention, with the combination group obtaining the highest form scores, followed by the all-pre-practice group, and finally the interspersed group. These findings suggest that several modeling exposures before practice and several more exposures in the early stages of practice were optimal for acquisition and retention of form.     

Fighting and playfighting in captive adolescent chimpanzees

Most young non-human primates spend a great deal of time playing alone or with conspecifics. Despite numerous studies on social play, its adaptive functions remain unclear. A longitudinal study comparing social interactions preceding and following rank reversals among four captive adolescent chimpanzees (Pan troglodytes schweinfurtii) suggests that social play during their adolescent period functioned in establishing and maintaining the dominance rank within dyads, without interventions from allies. Interventions were less frequent in rough-and-tumble play than in real fighting. Within dyads of competing chimpanzees, rough play was more frequent than gentle play. Both types of play were correlated with reciprocal aggression between competitors, and most play bouts terminated in truly aggressive conflict. Dominant individuals seemed to be responsible for maintaining play activity, by responding playfully to threats and aggression of contestants. In addition, dominant individuals more often initiated play encounters with their subordinates when the dominance relationships were clear, but subordinate individuals also initiated play sequences when they challenged the dominant's rank. The adaptive value and mechanisms of rough-and-tumble play are discussed in the context of adolescence. © 1994 Wiley-Liss, Inc.     

文化演化的认知视角——从个体社会学习出发探究文化动态性

文化演化是多学科共同关心的文化研究主题。文化演化的认知视角是心理学家切入该研究主题的一种方式, 它着眼于社会文化环境中的个体认知, 研究个体在社会学习过程中对文化信息的加工、改变、记忆与提取。文化演化的认知视角借用达尔文生物演化理论, 涉及文化传承、创新、选择三个子领域, 提出了文化演化的三原则: 遗传、变异、选择。文化传承的路径包括模仿和教导, 类型分为工具性与习俗性文化传承; 文化创新具有层次性, 人类特有的累积性文化演化建立在文化创新的基础之上, 体现了文化创新的代际传递; 基于行为生态学和人类认知机制的文化选择造成了文化信息的差异化适应。未来研究可以从研究概念、研究思路、研究方法三方面推进认知视角下的文化演化研究, 探索更高层次的文化创新, 扩展文化演化的前因变量, 结合新技术加深对文化演化的理解; 发挥文化演化对文化心理学研究的助推作用, 研究文化混搭、个性心理特征对文化演化的影响。     

THE CUMULATIVE QUALITY OF CULTURE EXPLAINS HUMAN UNIQUENESS

What explains the unique features of human culture? Culture is not uniquely human, but human culture is uniquely cumulative. Cumulative culture is a product of our collective intelligence and is supported by cognitive processes and learning strategies that enable people to acquire, transform, and transmit information and technologies within and across generations. Technological and social innovations are currently driving unprecedented changes in cultural complexity and diversity. Innovation is a cognitively and socially complex, multistep process that typically requires (cumulative) cultural learning to achieve. I argue that the technological solutions that characterize twenty-first-century innovation can only be explained by understanding both the capacity to learn from and build upon the insights of others and the transmission systems that store the knowledge and technologies of previous generations. Human uniqueness is a product of cumulative cultural learning, transmission, and innovation.     

Pedagogical cues encourage toddlers' transmission of recently demonstrated functions to unfamiliar adults

Young children use pedagogical cues as a signal that others' actions are social or cultural conventions. Here we show that children selectively transmit (enact in a new social situation) causal functions demonstrated pedagogically, even when they have learned and can produce alternative functions as well. Two‐year‐olds saw two novel toys, each with two functions. One experimenter demonstrated one function using pedagogical cues (eye contact and child‐directed speech) and a second experimenter demonstrated the alternative function using intentional actions towards the object, but without pedagogical cues. Children imitated both functions at equal rates initially, indicating equal causal learning from both types of demonstration. However, they were significantly more likely to enact the pedagogical function for a new adult not present during the initial demonstrations. These results indicate that pedagogical cues influence children's transmission of information, perhaps playing a role in the dissemination of cultural conventions from a young age.     

Social learning in captive African elephants (Loxodonta africana africana)

Social learning is a more efficient method of information acquisition and application than trial and error learning and is prevalent across a variety of animal taxa. Social learning is assumed to be important for elephants, but evidence in support of that claim is mostly anecdotal. Using a herd of six adult female African bush elephants (Loxodonta africana africana) at the San Diego Zoo’s Safari Park, we evaluated whether viewing a conspecific’s interactions facilitated learning of a novel task. The tasks used feeding apparatus that could be solved in one of two distinct ways. Contrary to our hypothesis, the method the demonstrating animal used did not predict the method used by the observer. However, we did find evidence of social learning: After watching the model, subjects spent a greater percentage of their time interacting with the apparatus than they did in unmodeled trials. These results suggest that the demonstrations of a model may increase the motivation of elephants to explore novel foraging tasks.