Early signs of language in cross-fostered chimpanzees

In cross-fostering the young of one species are reared by adults of another, as in the classical ethological studies of imprinting and song-learning. In our laboratory, infant chimpanzees were reared under human conditions that included two-way communication in American Sign Language (ASL) the gestural language of the deaf in North America. Here we describe the cross-fostering conditions of this laboratory and the contrast with operant conditioning. We also review the uses and the shapes of the signs in the vocabularies of five cross-fostered chimpanzees, tests demonstrating that the cross-fostered chimpanzees could use signs of ASL to communicate conceptual information to human observers whose only source of information was the signs of the chimpanzees, early development of inflections that resemble the early inflections of deaf human children, evidence based on errors and on inflections for duality of patterning, the continued use of sign language among the chimpanzees even when deprived of human input and the acquisition of signs by the infant Loulis from the cross-fostered chimpanzees.     

Ethological study of early language

Sign language studies of cross-fostered chimpanzees measure the effect of special rearing conditions on the development of very young chimpanzees. Cross-fostered chimpanzees, like human children, develop gradually in a process that takes many years. Here we discuss details of the procedure, the overlap between human and chimpanzee infants in the contents of the first 50-item vocabularies, and the ways in which the signs of the chimpanzees exhibit the fuzziness of natural language categories. We also compare the cross-fostering approach with more traditional modular approaches to the study of language-like behavior in nonhuman animals. Project Washoe was originally supported by grants MH-12154 from the National Institute of Mental Health and GB-7432 from the National Science Foundation. We gratefully acknowledge this support and the support that later sign language studies of chimpanzees have received since then from NIH, NSF, the National Geographic Society, the Grant Foundation, the Spencer Foundation, the University of Nevada, and the UNR Foundation.     

The role of cross-fostering in sign language studies of chimpanzees

In cross-fostering, the young of one species are reared by adults of another. In our cross-fostering laboratory, two-way communication by means of American Sign Language (ASL) brought the rearing conditions for chimpanzees much closer to those of human children than was possible in earlier studies. At the same time, ASL provided a means by which chimpanzees could express their intelligence in ways that permit closer comparisons with human children. Cross-fostered chimpanzees can communicate with human observers whose only source of information is the American Sign Language (ASL) signs of the chimpanzees. In order to show that the chimpanzees could name natural language categories — that the sign DOG could refer to any dog, FLOWER to any flower, SHOE to any shoe — each test trial was a first trial in that tests slides were presented only once. Analysis of errors showed that two aspects of the signs, gestural form and conceptual category, governed the distribution of errors. Like human adults and human children who sign, the chimpanzees modulated their signs in meaningful ways. Observations in field notes, video tape records, and systematic experiments illustrate how these modulations were related to the verbal and nonverbal, context and how they made signs more visible, more versatile, and more informative.     

Prelinguistic development of children and chimpanzees

What was unique about our laboratory was the detailed simulation of a human rearing environment for infant chimpanzees, together with continuous, intensive observation and recording of behavioral development. Communication in American Sign Language (ASL) was integrated into this procedure in a way that simulates the teaching of vocal and gestural languages to human children. The result is a comprehensive longitudinal record of the stage by stage development of two-way communication in sign language and its relation to basic aspects of behavioral development such as: locomotion, manipulative skills, observational learning, and social play. In the longitudinal records of five infant chimpanzees, we can trace the patterns of development and also examine the consistency of individual differences through the first five years. The central question is the relation between developing skills in the use of American Sign Language and the rest of behavioral development. To analyze factors that influence development, we have compared records for infant chimpanzees reared in the crossfostering laboratory with records for chimpanzees reared in the wild by their own mothers and chimpanzees reared in cages under conventional laboratory conditions, as well as with records of development for human children, reared in homes and reared in institutions.     

Chimpanzee vocal signaling points to a multimodal origin of human language

The evolutionary origin of human language and its neurobiological foundations has long been the object of intense scientific debate. Although a number of theories have been proposed, one particularly contentious model suggests that human language evolved from a manual gestural communication system in a common ape-human ancestor. Consistent with a gestural origins theory are data indicating that chimpanzees intentionally and referentially communicate via manual gestures, and the production of manual gestures, in conjunction with vocalizations, activates the chimpanzee Broca's area homologue--a region in the human brain that is critical for the planning and execution of language. However, it is not known if this activity observed in the chimpanzee Broca's area is the result of the chimpanzees producing manual communicative gestures, communicative sounds, or both. This information is critical for evaluating the theory that human language evolved from a strictly manual gestural system. To this end, we used positron emission tomography (PET) to examine the neural metabolic activity in the chimpanzee brain. We collected PET data in 4 subjects, all of whom produced manual communicative gestures. However, 2 of these subjects also produced so-called attention-getting vocalizations directed towards a human experimenter. Interestingly, only the two subjects that produced these attention-getting sounds showed greater mean metabolic activity in the Broca's area homologue as compared to a baseline scan. The two subjects that did not produce attention-getting sounds did not. These data contradict an exclusive "gestural origins" theory for they suggest that it is vocal signaling that selectively activates the Broca's area homologue in chimpanzees. In other words, the activity observed in the Broca's area homologue reflects the production of vocal signals by the chimpanzees, suggesting that this critical human language region was involved in vocal signaling in the common ancestor of both modern humans and chimpanzees.     

The ontogeny of human prosociality: behavioral experiments with children aged 3 to 8

Humans regularly engage in prosocial behavior that differs strikingly from that of even our closest living relatives, the chimpanzees (Pan troglodytes). In laboratory settings, chimpanzees are indifferent when given the opportunity to deliver valued rewards to conspecifics, while even very young human children have repeatedly been shown to behave prosocially. Although this broadly suggests that prosocial behavior in chimpanzees differs from that of young human children, the methods used in prior work with children have also differed from the methods used in studies of chimpanzees in potentially crucial ways. Here we test 92 pairs of 3–8-year-old children from urban American (Los Angeles, CA, USA) schools in a face-to-face task that closely parallels tasks used previously with chimpanzees. We found that children were more prosocial than chimpanzees have previously been in similar tasks, and our results suggest that this was driven more by a desire to provide benefits to others than a preference for egalitarian outcomes. We did not find consistent evidence that older children were more prosocial than younger children, implying that younger children behaved more prosocially in the current study than in previous studies in which participants were fully anonymous. These findings strongly suggest that humans are more prosocial than chimpanzees from an early age and that anonymity influences children's prosocial behavior, particularly at the youngest ages.     

Signs of intelligence in cross-fostered chimpanzees

In cross-fostering, the young of one species are reared by adults of another, as in the classical ethological studies of imprinting and song-learning. In our laboratory, infant chimpanzees were reared under human conditions that included two-way communication in American Sign Language (A.S.L.), the gestural language of the deaf in North America. A large body of evidence from five chimpanzees demonstrated stage by stage replication of basic aspects of the acquisition of speech and signs by hearing and deaf children. Here we review evidence that, under double-blind conditions: (i) the chimpanzees communicated information in A.S.L. to human observers; (ii) independent human observers agreed in their identification of the chimpanzee signs, (iii) the chimpanzees could use the signs to refer to natural language categories: DOG for any dog, FLOWER for any flower, SHOE for any shoe.     

Functional analysis of chimpanzee (Pan troglodytes) private signing

The chimpanzee's use of American Sign Language (ASL) to communicate with humans and with each other has been empirically demonstrated in several reports, but this is the first research to experimentally examine their use of sign language in a nonsocial fashion: private signing. This experiment examined the private signing behavior of five signing chimpanzees, using a remote videotaping technique with no human present. It was found that all five chimpanzees signed to themselves for a total of 368 instances. These instances of private signing were classified into nine different functional categories as has been done in the analysis of private speech and signing in hearing and deaf human children. Similar to humans, a few of the categories accounted for the majority of the instances of private signing. These findings empirically demonstrate a behavior similar to private speech and signing in humans.     

Multiple classification performance of juvenile chimpanzees,normal children,and retarded children

The cognitive capacities of juvenile chimpanzees, normal children, and retarded children were evaluated by using a nonverbal, Piagetian-type multiple classification task. The three groups of subjects were tested with the same two by two stimulus matrix, which was formed by combining two different colors with two different shapes. On each problem the subjects were required to select one of four stimulus items from the response tray and place it inside the empty cell of the stimulus matrix. It was found that the human and nonhuman primates tested were able to select objects with correct color and shape cues (C⁺/S⁺)significantly more often than the other objects with only color cues correct (C⁺/S^-), only shape cues correct (C^-/S⁺), or neither cue correct (C^-/S^-). Although the two groups of human children were able to select the C⁺/S⁺ objects about 100% of the time by the end of testing, the normal children required significantly fewer problems than the retarded children. The juvenile chimpanzees needed significantly more problems than the human subjects before they consistently selected the C⁺/S⁺ object on the first trial and attained a level of correct responding that was above chance. Moreover, their level of performance did not exceed 70% correct. These data suggest that the human children (both normal and retarded) used a conceptual strategy, while the juvenile chimpanzees employed a perceptual strategy to solve the multiple classification problems. The relationship between language and conceptual problem-solving strategies for Piagetiantype tasks is discussed. Portions of the data reported in this study were presented at the Annual Meeting of the Southern Psychological Association, Atlanta, Georgia, March 15–18, 1978.     

Communicative signaling activates 'Broca's' homolog in chimpanzees

Broca's area, a cerebral cortical area located in the inferior frontal gyrus (IFG) of the human brain, has been identified as one of several critical regions associated with the motor planning and execution of language. Anatomically, Broca's area is most often larger in the left hemisphere, and functional imaging studies in humans indicate significant left-lateralized patterns of activation during language-related tasks. If, and to what extent, nonhuman primates, particularly chimpanzees, possess a homologous region that is involved in the production of their own communicative signals remains unknown. Here, we show that portions of the IFG as well as other cortical and subcortical regions in chimpanzees are active during the production of communicative signals. These findings are the first to provide direct evidence of the neuroanatomical structures associated with the production of communicative behaviors in chimpanzees. Significant activation in the left IFG in conjunction with other cortical and subcortical brain areas during the production of communicative signals in chimpanzees suggests that the neurological substrates underlying language production in the human brain may have been present in the common ancestor of humans and chimpanzees.     

Children,but not chimpanzees,have facial correlates of determination

Facial expressions have long been proposed to be important agents in forming and maintaining cooperative interactions in social groups. Human beings are inordinately cooperative when compared with their closest-living relatives, the great apes, and hence one might expect species differences in facial expressivity in contexts in which cooperation could be advantageous. Here, human children and chimpanzees were given an identical task designed to induce an element of frustration (it was impossible to solve). In children, but not chimpanzees, facial expressions associated with effort and determination positively correlated with persistence at the task. By contrast, bodily indicators of stress (self-directed behaviour) negatively correlated with task persistence in chimpanzees. Thus, children exhibited more behaviour as they persisted, and chimpanzees exhibited less. The facial expressions produced by children, could, therefore, function to solicit prosocial assistance from others.     

Possible preadaptations to speech. A preliminary comparative

Human language is a unique phenomenon and its evolutionary origins are uncertain. In this paper we attempt to explore some of the preadaptations that might have contributed to the origin of human speech. The comparative approach we use is based on the assumption that all features of a species are functional, and that all features can be compared with those of other animals and correlated with certain lifestyles. Using this method we attempt to reconstruct the different evolutionary pathways of humans and chimpanzees after they split from a common ancestor. Previous results from comparative studies suggest human ancestors may not have evolved on the open African savannas as was once believed, but more probably were coastal omnivores feeding on plant matter and easy to catch invertebrates such as shellfish from beaches and shallow waters. Fossil and archaeological data suggest this coastal phase occurred at the beginning of the Pleistocene, whenHomo ergaster-erectus dispersed between East-Africa, North-Africa, South-Asia and Indonesia. This paper presents comparative data suggesting the various human speech skills may have had their origins at different times and may originally have had different functions. Possible preadaptations to speech include, for instance, musical skills present in a variety of primate species (sound production); airway closure and breath-hold diving for collecting seafood (voluntary breath control); and suction feeding adaptations for the consumption of fruit juice or certain seafoods (fine control of oropharyngeal movements). The different evolutionary pathways of chimpanzees and humans might explain why chimpanzees lack language skills and why human language is a relatively recent phenomenon.     

Imaginary Play in chimpanzees (Pan troglodytes)

Imaginary Play was studied in a group of five signing chimpanzees and it was found that chimpanzees engage in imaginary play similar to that found in human children. Fifteen hours of remote videotapes were analyzed for instances of imaginary play. Behaviors were defined as imaginary play by meeting a predetermined criteria which allowed them to be classified into one of six different categories of imaginary play. Six instances of imaginary play were found and these were classified into the two categories of Animation and Substitution. Observations of imaginary play in other research with chimpanzees were discussed.     

Behavioral responsiveness of young chimpanzees (Pan troglodytes) to a novel environment

Behavioral responsiveness to a novel environment was documented in 22 chimpanzees grouped according to age; 6-months, 1-year, 2-years and 5-years. An attachment figure, a human caretaker, accompanied each subject during the 15-min test sessions so as to preclude confounding of responses to novelty with separation responses. Extreme distress reported previously for chimpanzees and human children when tested alone in a novel situation was rarely observed in these tests when an attachment figure was present. Stereotyped rocking, an indication of mild distress occurred more frequently in the younger animals. Younger animals engaged in distal visual exploration of the environment while remaining close to the attachment figure, whereas the older animals locomoted more frequently and explored the environment directly with their hands. Repeated exposure to the environment reduced the differences among the 6-month, 1-year and 2-year groups. The 6-month group, however, continued to locomote least and least frequently engaged in tactile exploration. These data on chimpanzees resemble data on human children which suggest that an attachment figure: attenuates the distress exhibited by young individuals of these species when exposed to novel stimuli, and thereby provides a secure base which supports the exploration of novel stimuli, a prerequisite to behavioral adaptation.     

A structure-based repertoire of manual gestures in wild chimpanzees: statistical analyses of a graded communication system

Great ape gestural communication is considered important in understanding the evolution of human language as these share important features, namely, flexible and intentional signal use. Although gestural repertoires have been compiled for captive and wild primates, reports are largely qualitative. We quantify the morphological structure and variation of gestural signals identified in the repertoire of a community of wild chimpanzees. Gestures were classified on the basis of 29 morphological features, such as trajectory and orientation during the preparatory and stroke phases of a gesture. Hierarchical cluster and discriminant function analyses identified 30 morphologically distinct manual gesture types; the majority was subsequently correctly classified using a cross-validation technique, with incorrect classifications for rare gesture types only. Comparisons of this statistically determined repertoire with previous repertoires did not identify systematic variation between captive and wild chimpanzees. Moreover, consensus was not greater within studies of the same populations, highlighting the importance of systematic and well-documented inventories. Our morphologically based analyses indicate that manual gestures are best considered as graded rather than discrete communication signals, similar to some vocalisation systems. We discuss these findings in light of current theories of human language evolution.     

Social learning of a communicative signal in captive chimpanzees

The acquisition of linguistic competency from more experienced social partners is a fundamental aspect of human language. However, there is little evidence that non-human primates learn to use their vocalizations from social partners. Captive chimpanzees (Pan troglodytes) produce idiosyncratic vocal signals that are used intentionally to capture the attention of a human experimenter. Interestingly, not all apes produce these sounds, and it is unclear what factors explain this difference. We tested the hypothesis that these attention-getting (AG) sounds are socially learned via transmission between mothers and their offspring. We assessed 158 chimpanzees to determine if they produced AG sounds. A significant association was found between mother and offspring sound production. This association was attributable to individuals who were raised by their biological mother-as opposed to those raised by humans in a nursery environment. These data support the hypothesis that social learning plays a role in the acquisition and use of communicative vocal signals in chimpanzees.     

The development of contingent reciprocity in children

Cooperation between nonrelatives is common in humans. Reciprocal altruism is a plausible evolutionary mechanism for cooperation within unrelated pairs, as selection may favor individuals who selectively cooperate with those who have cooperated with them in the past. Reciprocity is often observed in humans, but there is only limited evidence of reciprocal altruism in other primate species, raising questions about the origins of human reciprocity. Here, we explore how reciprocity develops in a sample of American children ranging from 3 to 7.5 years of age, and also compare children's behavior to that of chimpanzees in prior studies to gain insight into the phylogeny of human reciprocity. Children show a marked tendency to respond contingently to both prosocial and selfish acts, patterns that have not been seen among chimpanzees in prior studies. Our results show that reciprocity increases markedly with age in this population of children, and by about 5.5 years of age children consistently match the previous behavior of their partners.     

Vocal responsiveness in male wild chimpanzees: implications for the evolution of language

Several captive chimpanzees and bonobos have learned to use symbols and to comprehend syntax. Thus, compared with other nonhumans, these animals appear to have unusual cognitive powers that can be recruited for communicative behavior. This raises the possibility that wild chimpanzee vocal communication is more complex than heretofore demonstrated. To examine this possibility, I investigated whether wild chimpanzee vocal exchanges exhibit uniquely human conversational attributes. The results indicate that wild chimpanzees vocalize at low rates, tend not to respond to calls that they hear, and, when they do respond, tend to give calls that are similar to the ones they have heard. Thus, chimpanzee vocal interactions resemble those of other primate species, and show no special similarity to human conversations. The results support the view that we need to explore cognitive and social continuities and discontinuities with nonhuman primates to understand the origin and evolution of language, but also emphasize the need for fine-grained analyses of wild chimpanzee vocal interactions.     

Wernicke's area homologue in chimpanzees (Pan troglodytes) and its relation to the appearance of modern human language

Human language is distinctive compared with the communication systems of other species. Yet, several questions concerning its emergence and evolution remain unresolved. As a means of evaluating the neuroanatomical changes relevant to language that accompanied divergence from the last common ancestor of chimpanzees, bonobos and humans, we defined the cytoarchitectonic boundaries of area Tpt, a component of Wernicke''s area, in 12 common chimpanzee brains and used design-based stereologic methods to estimate regional volumes, total neuron number and neuron density. In addition, we created a probabilistic map of the location of area Tpt in a template chimpanzee brain coordinate space. Our results show that chimpanzees display significant population-level leftward asymmetry of area Tpt in terms of neuron number, with volume asymmetry approaching significance. Furthermore, asymmetry in the number of neurons in area Tpt was positively correlated with asymmetry of neuron numbers in Brodmann''s area 45, a component of Broca''s frontal language region. Our findings support the conclusion that leftward asymmetry of Wernicke''s area originated prior to the appearance of modern human language and before our divergence from the last common ancestor. Moreover, this study provides the first evidence of covariance between asymmetry of anterior and posterior cortical regions that in humans are important to language and other higher order cognitive functions.     

Words matter: Reflections on language projects with chimpanzees and their implications

Beginning in the 1960s the first systematic projects dedicated to testing whether great apes could acquire some aspects of human language were conducted. The ape subjects demonstrated remarkable capacities to learn and use elements of either sign language or an artificial language. The results from research across several laboratories drew a mixture of excitement and skepticism, and critiques and debates have ensued since the earliest reports were published. This continues today. Terrace (2019, Nim: A chimpanzee who learned sign language. New York, NY: Columbia University Press) repeats many of the same points made decades earlier, and has added some additional critiques. That scientists hold different perspectives on what to conclude from ape language studies is expected. However, any conclusion one draws should be based upon available evidence, which we outline in this review. We also address the critiques offered by Terrace (2019), including the stance that apes cannot understand or use words. Focusing on symbol use by chimpanzees and bonobos we describe evidence that argues for understanding of words, including capacities for declarative communication and intersubjectivity found in these apes. We conclude that the many decades of research using a variety of symbol systems challenges the absolutist position that chimpanzees and bonobos cannot learn language or understand the concept of a word.