The effects of early experience on adult copulatory behavior in zoo-born chimpanzees (Pan troglodytes)

This paper examines the relationship between the early rearing experience of zoo-born, zoo-reared chimpanzees (Pan troglodytes) and the subsequent occurrence of successful copulation as adults. Developmental histories were acquired for 71 subjects via questionnaires and phone interviews. The following variables related to aspects of chimpanzees' early rearing experience were examined: (1) Rearing conditions, that is, hand reared alone, reared with siblings or peers, or reared by at least one adult conspecific. (2) Age removed from mother. (3) Sex of subject, and/or (4) participation in shows. Each of these variables was then compared to the subjects' sexual competence, defined here as having been observed to exhibit functional copulatory behavior as adults. Seventy-five percent of the subjects were observed to exhibit functional copulatory behavior on at least one occasion. No single component of rearing was successful in producing sexually competent adults 100% of the time; no rearing condition, as defined in this study, resulted in reproductive failure for all subjects experiencing those conditions. Chimpanzees that were hand reared alone, that is, in the total absence of conspecifics, were least likely to copulate as adults; about half of those chimpanzees that were reared with siblings or peers (and in the absence of adult conspecifics) copulated as adults. Almost 90% of those chimpanzees that were reared with at least one adult conspecific copulated as adults. Chimpanzees removed from their mothers at an early age (less than one year of age) were less likely to reproduce as adults. Male and female chimpanzees were equally likely to reproduce as adults. Participation in shows or demonstrations appeared to have no effect on chimpanzees' ability to copulate as adults. © 1994 Wiley-Liss, Inc.     

Effects of early rearing conditions on problem‐solving skill in captive male chimpanzees (Pan troglodytes)

Early rearing conditions of captive chimpanzees characterize behavioral differences in tool use, response to novelty, and sexual and maternal competence later in life. Restricted rearing conditions during early life hinder the acquisition and execution of such behaviors, which characterize the daily life of animals. This study examined whether rearing conditions affect adult male chimpanzees' behavior skills used for solving a problem with acquired locomotion behavior. Subjects were 13 male residents of the Chimpanzee Sanctuary Uto: 5 wild‐born and 8 captive‐born. A pretest assessed bed building and tool use abilities to verify behavioral differences between wild‐ and captive‐born subjects, as earlier reports have described. Second, a banana‐access test was conducted to investigate the problem‐solving ability of climbing a bamboo pillar for accessing a banana, which might be the most efficient food access strategy for this setting. The test was repeated in a social setting. Results show that wild‐born subjects were better able than captive‐born subjects to use the provided materials for bed building and tool use. Results of the banana‐access test show that wild‐born subjects more frequently used a bamboo pillar for obtaining a banana with an efficient strategy than captive‐born subjects did. Of the eight captive‐born subjects, six avoided the bamboo pillars to get a banana and instead used, sometimes in a roundabout way, an iron pillar or fence. Results consistently underscored the adaptive and sophisticated skills of wild‐born male chimpanzees in problem‐solving tasks. The rearing conditions affected both the behavior acquisition and the execution of behaviors that had already been acquired. Am. J. Primatol. 72:626–633, 2010. © 2010 Wiley‐Liss, Inc.     

Variables influencing the origins of diverse abnormal behaviors in a large sample of captive chimpanzees (Pan troglodytes)

The developmental origin of abnormal behaviors is generally associated with early rearing environments that lack sufficient physical and sensory stimulation. However, other factors should also be considered. A large sample of captive chimpanzees (128 males and 140 females) was surveyed for the presence or absence of 18 abnormal behaviors. Origin variables included the subject's source (zoo, pet, performer, or laboratory), rearing (mother‐ or hand‐reared), and sex. Animals were assessed while held at the Primate Foundation of Arizona, University of Texas M. D. Anderson Cancer Center, or White Sands Research Center. There was a confound among origin variables; more hand‐reared animals than expected were from laboratories. Logistic regression tested the relationship of rearing and source, with sex as a secondary predictor variable, to each of the abnormal behaviors. There was no clear association between any abnormal behavior and source. However, for coprophagy, relative to animals from the laboratory, zoo animals tended to show a higher prevalence, while performers tended to show a lower prevalence (when rearing and sex were controlled). Rocking and self‐sucking were significantly more likely in hand‐reared animals. Coprophagy and depilation of self were significantly more likely in mother‐reared animals. When rearing and source were statistically controlled, the only significant sex difference was a higher prevalence of coprophagy in females and a higher prevalence of rocking in males. In a second, smaller sample of 25 males and 33 females from Southwest Foundation for Biomedical Research, no significant sex association was found for coprophagy, urophagy, rocking, or self‐depilation. In this second sample, coprophagy was also significantly more likely in mother‐reared than hand‐reared subjects. The association of some abnormal behaviors with mother‐rearing suggests that some form of social learning may be involved in the origin of some of these behavior patterns. This indicates that some abnormal behaviors may not be always be indicative of reduced psychological well‐being in captive chimpanzees. Am. J. Primatol. 48:15–29, 1999. © 1999 Wiley‐Liss, Inc.     

Controllability in environmental enrichment for captive chimpanzees (Pan troglodytes)

This study considers the use of nonsocial environmental enrichment by captive chimpanzees at the Primate Foundation of Arizona. The goal was to determine whether a relationship existed between controllability of enrichment items by captive chimpanzees and frequency of use. The study measured controllability, the ability of nonhuman animals to alter aspects of their environment by the potential destructibility of the enrichment item. This study examined additional factors that may affect enrichment use: individual age, sex, rearing history, social group composition, and availability of outdoor access. The chimpanzees in the study used destructible items—the enrichment category with the highest level of controllability—more than indestructible items across all age, sex, and rearing classes. Thus, controllability seems to be an important factor in chimpanzee enrichment. Younger individuals and groups with outdoor access used enrichment more than did older individuals and groups with indoor-only access. Individual sex, rearing history, and social group composition had minimal effects on enrichment use. These results support the importance of control to captive chimpanzees and further enable captive management to customize enrichment programs to the needs of particular animals.     

Pestle-pounding behavior of wild chimpanzees at Bossou,Guinea: A newly observed tool-using behavior

A new type of tool-using behavior was observed in a group of wild chimpanzees (Pan troglodytes verus) at Bossou, Guinea. The chimpanzees used the leaf-petiole of oil-palm trees (Elaeis guineensis) as a pounding tool to deepen a hole in the oil-palm crown which appeared after the chimpanzees had pulled out the central young shoots. Finally, the chimpanzees extracted and ate the apical meristem or apical bud of the oil-palm tree which is edible but inaccessible without such tool use. The motor pattern which the chimpanzees employed is similar to that used for termite-nest digging but it is more exaggerated and requires great force. The behavior is reminiscent of pestlepounding. The chimpanzees exploit substantial amounts of food with this tool-using skill, compensating for insufficient fruit foods in the primary forest. This tool-using behavior was first observed in 1990 and, to date, almost half of the group members have been confirmed to use the pestle tool. It appears that this tool-using behavior was invented recently and has since spread widely throughout the group as a habitual one.     

Genetic basis in motor skill and hand preference for tool use in chimpanzees (Pan troglodytes)

Chimpanzees are well known for their tool using abilities. Numerous studies have documented variability in tool use among chimpanzees and the role that social learning and other factors play in their development. There are also findings on hand use in both captive and wild chimpanzees; however, less understood are the potential roles of genetic and non-genetic mechanisms in determining individual differences in tool use skill and laterality. Here, we examined heritability in tool use skill and handedness for a probing task in a sample of 243 captive chimpanzees. Quantitative genetic analysis, based on the extant pedigrees, showed that overall both tool use skill and handedness were significantly heritable. Significant heritability in motor skill was evident in two genetically distinct populations of apes, and between two cohorts that received different early social rearing experiences. We further found that motor skill decreased with age and that males were more commonly left-handed than females. Collectively, these data suggest that though non-genetic factors do influence tool use performance and handedness in chimpanzees, genetic factors also play a significant role, as has been reported in humans.     

Do Chimpanzees Use Weight to Select Hammer Tools?

The extent to which tool-using animals take into account relevant task parameters is poorly understood. Nut cracking is one of the most complex forms of tool use, the choice of an adequate hammer being a critical aspect in success. Several properties make a hammer suitable for nut cracking, with weight being a key factor in determining the impact of a strike; in general, the greater the weight the fewer strikes required. This study experimentally investigated whether chimpanzees are able to encode the relevance of weight as a property of hammers to crack open nuts. By presenting chimpanzees with three hammers that differed solely in weight, we assessed their ability to relate the weight of the different tools with their effectiveness and thus select the most effective one(s). Our results show that chimpanzees use weight alone in selecting tools to crack open nuts and that experience clearly affects the subjects' attentiveness to the tool properties that are relevant for the task at hand. Chimpanzees can encode the requirements that a nut-cracking tool should meet (in terms of weight) to be effective.     

Development of the visual preference of chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>) for photographs of primates: effect of social experience

In a study by Tanaka (2003) five captive chimpanzees preferred photographs of humans to those of chimpanzees. All the subjects were raised by humans and lived in captivity for many years. This suggests their preference might have developed through social experience. In this study examined this hypothesis by using three young chimpanzees raised by their mothers in a captive chimpanzee community. The young chimpanzees were tested four times before six years of age. I also tested eight adult chimpanzees that had been in captivity for more than 20 years. Each subject was presented with digitized color photographs of different species of primates on a touch-sensitive screen. The subjects received a food reward when they touched a photograph, irrespective of which photograph they touched. All the adult chimpanzees touched photographs of humans more frequently than those of any other species of primate. Two of the young chimpanzees showed no species preference before reaching 5 years of age, when they started to show preference for humans. The remaining young chimpanzee consistently preferred chimpanzees. These results suggest that development of visual preference of chimpanzees is affected by social experience during infancy.     

Tool use by captive chimpanzees at an artificial termite mound

This study examines tool use by a colony of captive chimpanzees at an artificial termite mound. The mound, constructed of concrete, simulates the termite mounds which are used as food sources by wild chimpanzees who extract the termites using grass or twig-type tools. In the present study, tool availability was manipulated, specifically the type of tool, and the distance of tool material from the mound. The type of food available in the mound was also varied. Tool-making and tool-using behavior was examined in relation to individual, age, and sex differences. The artificial mound proved to be a viable simulation of the naturally occurring mounds, with most of the chimpanzees exploiting the food in the mound by using tools over the period of study. Interesting individual differences emerged in the way that the chimpanzees selected and used tools, some preferring to move some distance from the mound to collect “off-the-peg” tools, others preferring to sit and fashion a tool from material available nearer the mound. Also, some chimpanzees used both ends of a tool, while others used only one end. There were significant age differences in activity at the mound, the younger chimpanzees spending more time at the mound, using tools previously used by others, and manipulating the mound holes manually. Sex differences, although not significant, were apparent. The artificial mound provides the chimpanzees with a stimulating and rewarding activity, interest and enjoyment for the public, and an opportunity for researchers to study tool use under more controlled conditions than are possible in the field.     

Rearing environment influences boldness and prey acquisition behavior,and brain and lens development of bull trout

Animals reared in barren captive environments exhibit different developmental trajectories and behaviors than wild counterparts. Hence, the captive phenotypes may influence the success of reintroduction and recovery programs for threatened and endangered species. We collected wild bull trout embryos from the Metolius River Basin, Oregon and reared them in differing environments to better understand how captivity affects the bull trout Salvelinus confluentus phenotype. We compared the boldness and prey acquisition behaviors and development of the brain and eye lens of bull trout reared in conventional barren and more structurally complex captive environments with that of wild fish. Wild fish and captive reared fish from complex habitats exhibited a greater level of boldness and prey acquisition ability, than fish reared in conventional captive environments. In addition, the eye lens of conventionally reared bull trout was larger than complex reared captive fish or same age wild fish. Interestingly, we detected wild fish had a smaller relative cerebellum than either captive reared treatment. Our results suggest that rearing fish in more complex captive environments can create a more wild-like phenotype than conventional rearing practices. A better understanding of the effects of captivity on the development and behavior of bull trout can inform rearing and reintroduction programs though prediction of the performance of released individuals.     

Comparing the performances of apes (Gorilla gorilla, Pan troglodytes, Pongo pygmaeus) and human children (Homo sapiens) in the floating peanut task

Recently, Mendes et al. [1] described the use of a liquid tool (water) in captive orangutans. Here, we tested chimpanzees and gorillas for the first time with the same "floating peanut task." None of the subjects solved the task. In order to better understand the cognitive demands of the task, we further tested other populations of chimpanzees and orangutans with the variation of the peanut initially floating or not. Twenty percent of the chimpanzees but none of the orangutans were successful. Additional controls revealed that successful subjects added water only if it was necessary to obtain the nut. Another experiment was conducted to investigate the reason for the differences in performance between the unsuccessful (Experiment 1) and the successful (Experiment 2) chimpanzee populations. We found suggestive evidence for the view that functional fixedness might have impaired the chimpanzees' strategies in the first experiment. Finally, we tested how human children of different age classes perform in an analogous experimental setting. Within the oldest group (8 years), 58 percent of the children solved the problem, whereas in the youngest group (4 years), only 8 percent were able to find the solution.     

Manual Laterality for Simple Reaching and Bimanual Coordinated Task in Naturalistic Housed <Emphasis Type="Italic">Pan troglodytes</Emphasis>

Researchers have shown renewed interest in the study of manual lateralization in chimpanzees. Currently there is no consensus confirming the presence or absence of manual dominance at a species level, mainly for populations in the wild and in semicaptivity. We aimed to evaluate the manual laterality in a group of chimpanzees in an intermediate setting (semicaptivity) via 2 tasks: one simple and unimanual (simple reaching) and the other complex and bimanual (tube task). We replicated the same experiments from Hopkins in a new and different sample of chimpanzees. In simple reaching, the hand is used to gather food and the type of grip and the posture are evaluated. The tube task assesses the hand used to extract food from the tube and the method of extraction (digital or instrumental). Through the handedness index we observed that the subjects show clear and strong individual preferences for both tasks (100% lateralized subjects in the tube task; 86% in simple reaching), although we did not detect population preferences for any of the tasks. However, considering both tasks jointly (multiple evaluation), it was possible to detect, for the first time, skilled manual dominance at a group level in semicaptive chimpanzees in one multitask index and borderline significance in a second multitask index.     

Population-Level Right Handedness for a Coordinated Bimanual Task in Chimpanzees: Replication and Extension in a Second Colony of Apes

The purpose of this study was to evaluate the reliability of previously published findings on hand preferences in chimpanzees by evaluating hand use in a second colony of captive chimpanzees. We assessed hand preferences for a coordinated bimanual task in 116 chimpanzees (Pan troglodytes) at the University of Texas M. D. Anderson Cancer Center and compared them to previously published findings in captive chimpanzees at the Yerkes National Primate Research Center. The new sample showed significant population-level right handedness, which is consistent with previously published findings in the Yerkes chimpanzees. Combined data on the 2 chimpanzee colonies, revealed a significant effect of rearing history on hand preference, with wild-caught chimpanzees showing less right-handedness than captive-born mother-reared chimpanzees. We discuss the results in terms of the role of early environment on the development of laterality.     

Baboons, chimpanzees, and tools

Wild chimpanzees make and use tools with far greater frequency and variety than wild baboons. Sampling differences; differences in the sensory, motor, or cognitive capacities for skilled tool use; or environmental differences are not responsible. Fortuitous discovery of novel forms of tool behavior probably occurs as rarely among chimpanzees as among baboons. However, research on wild and captive chimpanzees reveals that the discovery is transmitted quickly among chimpanzee groups by observation learning, and thus becomes well-established in the group's behavioral repertoire. In contrast, study of captive baboons reveals that they acquire little information about tool behavior by observation. With little observation learning the behavior is not likely to be acquired by other group members since repeated independent discovery is improbable. Thus chimpanzees' more proficient tool behavior is due mainly to their greater capacity for observation learning. This suggests that the advent of hominid tool traditions was accompanied by a capacity for facile observation learning and thus, perhaps, by language ability.     

The use of tools by primates: definition, criteria and hypotheses of its appearance in anthropogenesis

The different approaches to the definition of "tool-using" and "tool" are discussed. The definitions of tool-using are given, the hypotheses of the tool-using origin in phylogenesis of primates: emotional, social tradition, playing, accumulations' objects are adduced. The modern data on tool-using for catching ants and termites, for cracking palm-nuts by wild chimpanzees under natural inhabitation, population and sex differences in tool-using are given. A conclusion has been made that tool-using in apes is qualitatively new form of behaviour arising in phylogenesis of primates and demonstrating us prerequisity of tool-using of early hominids.     

Acquisition of a socially learned tool use sequence in chimpanzees: Implications for cumulative culture

Cumulative culture underpins humanity's enormous success as a species. Claims that other animals are incapable of cultural ratcheting are prevalent, but are founded on just a handful of empirical studies. Whether cumulative culture is unique to humans thus remains a controversial and understudied question that has far-reaching implications for our understanding of the evolution of this phenomenon. We investigated whether one of human's two closest living primate relatives, chimpanzees, are capable of a degree of cultural ratcheting by exposing captive populations to a novel juice extraction task. We found that groups (N = 3) seeded with a model trained to perform a tool modification that built upon simpler, unmodified tool use developed the seeded tool method that allowed greater juice returns than achieved by groups not exposed to a trained model (non-seeded controls; N = 3). One non-seeded group also discovered the behavioral sequence, either by coupling asocial and social learning or by repeated invention. This behavioral sequence was found to be beyond what an additional control sample of chimpanzees (N = 1 group) could discover for themselves without a competent model and lacking experience with simpler, unmodified tool behaviors. Five chimpanzees tested individually with no social information, but with experience of simple unmodified tool use, invented part, but not all, of the behavioral sequence. Our findings indicate that (i) social learning facilitated the propagation of the model-demonstrated tool modification technique, (ii) experience with simple tool behaviors may facilitate individual discovery of more complex tool manipulations, and (iii) a subset of individuals were capable of learning relatively complex behaviors either by learning asocially and socially or by repeated invention over time. That chimpanzees learn increasingly complex behaviors through social and asocial learning suggests that humans' extraordinary ability to do so was built on such prior foundations.     

Tool-using and -making behavior in wild chimpanzees at Bossou,Guinea

The behavior of wild chimpanzees at Bossou, Guinea, was studied from November 1976 to May 1977 recognizing each chimpanzee without artificial feeding. During the study period some tool-using and tool-making behavior was observed, as follows: (1) Although water drinking using a “leaf-sponge” was not seen, that using a “leaf-spoon” was observed for taking water from the hollow of a tree. (2) “Termite fishing” was not seen in this group although there were many termite hills in the moving range of the chimpanzees. They dug termites from the hollow of a tree by pounding with a small stick. Similar use of a stick was made for digging up the resin from a tree. (3) “Aimed throwing” was frequently observed in adult males for attacking an observer, and in adolescents and juveniles as mischief against an observer or for their own play. (4) “Nut cracking” with a pair of stones was seen for removing the ovule from palm-seeds. Particular stones were repeatedly used by many chimpanzees for a long period. (5) “Branch hauling” represented difficult work. Patient and inventive manufacture of proper sticks was necessary for capturing branches which they were unable to reach normally. Local variations in the tool-using patterns and manufacturing ability of chimpanzees are discussed.     

Discovering and learning tool-use for fishing honey by captive chimpanzees

Wild chimpanzees commonly use sticks to fish for termites, ants or honey. This ability seems to be socially transmitted to juveniles by their mothers across generations. In a natural environment, the limited visibility of this behavior with regards to the extent of stick's insertion and about the success or failure in fishing hinders the study of the underlying learning processes. This study explores the discovery and learning of tool use for fishing honey in an artificial hive by a group of four captive chimpanzees. The discovery of tool use was accidental and coactive. The speed with which the group of experimentally naive chimpanzees discovered and learned tool use suggests that wild chimpanzees of different populations independently discovered the fishing behavior. The alpha male and his ally learned before the subordinates. Here, trial-and-error learning was, as in monkeys, the main process allowing the acquisition of the tool-use technique. However, the observation of conspecifics allowed the orientation of the experimentation by the selection of clues. As suggested by Tomaselloet al. (1987). it is the understanding of the function of the tool,i.e. the cause-effect relations between the action of the demonstrator, the type of tool and the task to accomplish which confer to chimpanzees and advantage over monkeys.     

Complex tool sets for honey extraction among chimpanzees in Loango National Park, Gabon

Homo faber was once proposed as a label for humans specifically to highlight their unique propensity for tool use. However, new observations on complex tool use by the chimpanzees of Loango National Park, Gabon, expand our knowledge about tool-using abilities in Pan troglodytes. Chimpanzees in Loango, when using tools to extract honey from three types of bee nests, were observed to regularly use three- to five-element tool sets. In other words, different types of tools were used sequentially to access a single food source. Such tool sets included multi-function tools that present typical wear for two distinct uses. In addition, chimpanzees exploited underground bee nests and used ground-perforating tools to locate nest chambers that were not visible from the ground surface. These new observations concur with others from Central African chimpanzees to highlight the importance of honey extraction in arguments favoring the emergence of complex tool use in hominoids, including different tool types, expanded tool sets, multifunction tools, and the exploitation of underground resources. This last technique requires sophisticated cognitive abilities concerning unseen objects. A sequential analysis reveals a higher level of complexity in honey extraction than previously proposed for nut cracking or hunting tools, and compares with some technologies attributed to early hominins from the Early and Middle Stone Age. A better understanding of similarities in human and chimpanzee tool use will allow for a greater understanding of tool-using skills that are uniquely human.     

Do chimpanzees cooperate in a learning task?

The aim of this study was to assess the ability of chimpanzees (Pan troglodytes) to cooperate in an instrumental task. A specially constructed fruit distributor was presented to a group of six captive chimpanzees. A cooperative response required two chimpanzees: both had to pull a handle simultaneously to make a fruit fall into the cage. The dominant male of the group and an infant produced most of the operant responses, and the male got nearly all the fruits. Other conspecifics avoided the dominant male at the apparatus. Social influences appear to limit the possibility of co-operation between individuals because a certain level of interindividual tolerance is required. The results revealed a significant increase in the number of pulls each time both chimpanzees were together at the apparatus. Operant chimpanzees learn to coordinate their actions in time and space.