Preliminary assessment of methods used to demonstrate nut-cracking behavior to five captive chimpanzees (Pan troglodytes)

Chimpanzees acquire nut-cracking skills by observation and trial and error. Studies of captive chimpanzees have shown the effectiveness of a skilled demonstrator. We examined the effectiveness of 3 live demonstration forms from which subjects could learn nut-cracking skills: a video of proficient conspecifics, human demonstration and the presence of a skilled conspecific performing the task. A male subject did not learn to crack open nuts after viewing a video of proficient conspecifics but quickly learned the skill following a demonstration by a human facilitator. Subsequently, 4 female chimpanzees were given the opportunity to learn the skill from the now proficient male, as well as from a video and human demonstration, but failed to do so.     

‘Ghost’ experiments and the dissection of social learning in humans and animals

The focus of this review is the experimental techniques used to identify forms of social learning shown by humans and nonhuman animals. Specifically, the ‘ghost display’ and ‘end‐state’ conditions, which have been used to tease apart imitative and emulative learning are evaluated. In a ghost display, the movements of an apparatus are demonstrated, often through the discrete use of fishing‐line or hidden mechanisms, without a live model acting directly upon the apparatus so that the apparatus appears to be operated as if by a ‘ghostly’ agent. In an end‐state condition, an observing individual is shown the initial state of the test apparatus, the apparatus is then manipulated out‐of‐sight and then represented to the individual in its final state. The aim of the ghost display condition is to determine whether individuals are able to emulate by replicating the movements of an apparatus, or perform a task, without requiring information about the bodily movements required to do so (imitation). The end‐state condition is used to identify goal‐emulation by assessing whether the observer can replicate the steps required to solve the task without having been shown the required body actions or task movements. The responses of individuals tested with either the ghost display and/or end‐state conditions are compared to those of further individuals who have observed a full demonstration by either a human experimenter or a conspecific. The responses of a control group, to whom no information has been provided about the test apparatus or required actions, are also compared and evaluated. The efficacy of these experimental techniques employed with humans, nonhuman primates, dogs, rats and birds are discussed and evaluated. The experiments reviewed herein emphasise the need to provide ghost displays and end‐state conditions in combination, along with full live demonstrations and a no‐information control. Future research directions are proposed.     

Observational learning in chimpanzees and children studied through 'ghost' conditions

Emulation has been distinguished from imitation as a form of observational learning because it focuses not on the model's actions but on the action's environmental results. Whether a species emulates, imitates or displays only simpler observational learning is expected to have profound implications for its capacity for cultural transmission. Chimpanzees' observational learning has been suggested to be primarily emulative, but this is an inference largely based upon low fidelity copying in experiments when comparing chimpanzees with humans rather than direct testing. Here we test directly for emulation learning by chimpanzees and children using a 'ghost' condition in which a sliding door obscuring a reward was moved to left or right with no agent visible, a context associated with the only published evidence for emulation learning in a non-human species (pigeons). Both children and chimpanzees matched the observed direction of ghost door movement on their first test trial. This is the first evidence for emulation in a non-human primate in the restricted context of a ghost condition. However, only the children continued to match in later trials. Individuals of both species continued to match with 99% or better fidelity when viewing a conspecific model operates the door. We conclude that chimpanzees can and will display emulation learning when the task is as simple as the present one, which contrasts with a failure to do so in a more complex manipulative task tested earlier. However, even with a simple task, emulation alone creates only fleeting fidelity compared with the opportunity to copy a conspecific, when considerable conformity is displayed.     

A potent effect of observational learning on chimpanzee tool construction

Although tool use occurs in diverse species, its complexity may mark an important distinction between humans and other animals. Chimpanzee tool use has many similarities to that seen in humans, yet evidence of the cumulatively complex and constructive technologies common in human populations remains absent in free-ranging chimpanzees. Here we provide the first evidence that chimpanzees have a latent capacity to socially learn to construct a composite tool. Fifty chimpanzees were assigned to one of five demonstration conditions that varied in the amount and type of information available in video footage of a conspecific. Chimpanzees exposed to complete footage of a chimpanzee combining the two components to retrieve a reward learned to combine the tools significantly more than those exposed to more restricted information. In a follow-up test, chimpanzees that constructed tools after watching the complete demonstration tended to do so even when the reward was within reach of the unmodified components, whereas those that spontaneously solved the task (without seeing the modification process) combined only when necessary. Social learning, therefore, had a powerful effect in instilling a marked persistence in the use of a complex technique at the cost of efficiency, inhibiting insightful tool use.     

A note on the responses of chimpanzees (Pan troglodytes) to live self-images on television monitors

The majority of studies on self-recognition in animals have been conducted using a mirror as the test device; little is known, however, about the responses of non-human primates toward their own images in media other than mirrors. This study provides preliminary data on the reactions of 10 chimpanzees to live self-images projected on two television monitors, each connected to a different video camera. Chimpanzees could see live images of their own faces, which were approximately life-sized, on one monitor. On the other monitor, they could see live images of their whole body, which were approximately one-fifth life-size, viewed diagonally from behind. In addition, several objects were introduced into the test situation. Out of 10 chimpanzees tested, 2 individuals performed self-exploratory behaviors while watching their own images on the monitors. One of these two chimpanzees successively picked up two of the provided objects in front of a monitor, and watched the images of these objects on the monitor. The results indicate that these chimpanzees were able to immediately recognize live images of themselves or objects on the monitors, even though several features of these images differed from those of their previous experience with mirrors.     

Interactive video playback and opponent assessment in lizards

Video playback has been used to explore many issues in animal communication, but the scope of this work has been constrained by the lack of stimulus-subject interaction. In many natural contexts, each participant's signalling behaviour is dependent from moment-to-moment on that of the other. Analyses of acoustic communication demonstrate the value of reproducing such social contingencies. We assessed the utility of interactive playback for studies of visual signalling by comparing the responses of male Jacky dragons, Amphibolurus muricatus, to interactive and non-interactive digital video playbacks of a life-sized conspecific. Displays produced by lizards in the interactive condition had the effect of suppressing the aggressive display of their simulated opponent. Each stimulus sequence generated during an interactive playback was subsequently played to a size-matched control animal. Males that could interact with the video stimulus responded principally with aggressive displays, while those that could not produced a mixture of aggressive and appeasement signals. Adding a degree of receiver responsiveness is hence sufficient to alter the type of signal evoked, even when video stimuli are physically identical. Interactive playback permits the experimental study of a broader range of theoretical topics and can enhance the realism of video stimuli.     

Digital video playback and visual communication in lizards

Experimental analyses of dynamic visual signals have to overcome the technical obstacle of reproducing complex motor patterns such as those found in courtship and threat displays. Video playback offers a potential solution to this problem, but it has recently been criticized because of sensory differences between humans and nonhuman animals, which suggest that video stimuli might be perceived as deficient relative to live conspecifics. Quantitative comparisons are therefore necessary to determine whether video sequences reliably evoke natural responses. Male Jacky dragons, Amphibolurus muricatus, compete for territories using complex displays delivered in a rapid stereotyped sequence. We evaluated video playback as a technique for studying this visual signal. Digital video sequences depicting a life-sized displaying male were indistinguishable from live male conspecifics in the rate and structure of aggressive displays evoked. Other measures of social behaviour suggested that video stimuli were more effective in this context. Lizards produced significantly more appeasement displays and had higher rates of substrate licking and locomotor activity in response to video playback than to confined male opponents, which failed to produce aggressive displays. Lizards tracked temporal changes in the display rate of video stimuli and were also sensitive to individual differences in morphology and behaviour between video exemplars. These results show that video stimuli are appropriate for the experimental analysis of Jacky dragon aggressive displays. We compare the potential shortcomings of video playback with those of other techniques and conclude that no approach offers a panacea, but that several have complementary characteristics. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Considerations in video playback design: using optic flow analysis to examine motion characteristics of live and computer-generated animation sequences

The increasing use of the video playback technique in behavioural ecology reveals a growing need to ensure better control of the visual stimuli that focal animals experience. Technological advances now allow researchers to develop computer-generated animations instead of using video sequences of live-acting demonstrators. However, care must be taken to match the motion characteristics (speed and velocity) of the animation to the original video source. Here, we presented a tool based on the use of an optic flow analysis program to measure the resemblance of motion characteristics of computer-generated animations compared to videos of live-acting animals. We examined three distinct displays (tail-flick (TF), push-up body rock (PUBR), and slow arm wave (SAW)) exhibited by animations of Jacky dragons (Amphibolurus muricatus) that were compared to the original video sequences of live lizards. We found no significant differences between the motion characteristics of videos and animations across all three displays. Our results showed that our animations are similar the speed and velocity features of each display. Researchers need to ensure that similar motion characteristics in animation and video stimuli are represented, and this feature is a critical component in the future success of the video playback technique.     

Response to Video and Computer-animated Images by the Tiger Barb, Puntius tetrazona

Video playback has become an important tool for testing certain questions about animal visual perception. While a few studies have compared the response of test subjects to live and video stimuli, it is generally assumed a priori that the animals will respond to the video image as if it were real. Since video devices are tuned to the spectral sensitivities and flicker fusion frequencies of the human eye and brain, it is important to conduct preliminary tests to compare the response of test subjects between live and video images. Here we compare schooling behavior of tiger barbs, Puntius tetrazona, when presented with a school of live fish, an analog video school of fish and a video of a computer-animated school of fish. Test subjects did not show a significantly different schooling behavior response to any of these stimuli. Additionally, when given the choice of schooling with a conspecific versus a heterospecific, tiger barbs schooled significantly more often with the conspecifics for all three stimulus formats. This study provides evidence that tiger barbs do not choose to discriminate between live and video models of fish and that this technique can be used to test future questions concerning visual perception for this species.     

Differential sensitivity to conspecific and allospecific cues in chimpanzees and humans: a comparative eye-tracking study

Previous studies have shown that a variety of animals including humans are sensitive to social cues from others and shift their attention to the same objects attended to by others. However, little is known about how animals process conspecifics'' and another species'' actions, although primates recognize conspecific faces better than those of another species. In this study, using unrestrained eye-tracking techniques, we first demonstrated that conspecific social cues modulated looking behaviours of chimpanzees more than human cues, whereas human observers were equally sensitive to both species. Additionally, first pass gaze duration at the face indicates that chimpanzees looked at the chimpanzees'' face longer than the human face, suggesting that chimpanzees might extract more referential information from a conspecific face. These results also imply that a unique ability for extracting referential information from a variety of social objects has emerged during human evolution.     

实体、镜像及影像对ICR小鼠取食行为的影响

镜像、影像等虚拟条件可能影响动物的觅食行为和活动时间分配。利用控制性围栏实验观察了同种个体实体、镜像及影像对ICR小鼠(Mus musculus)的觅食行为和活动时间分配的影响。发现实验鼠与实体接触时间显著高于与镜像、影像的接触时间;实体存在时,实验鼠显著增加了在低竞争风险区域贮藏食物量和巢外活动时间;影像存在时,实验鼠显著减少了巢外活动时间。结果提示ICR小鼠在实体、镜像、影像条件下表现出不同的觅食策略和活动时间分配,其觅食行为具有一定的可塑性。该结果可能为进一步研究ICR小鼠的行为及饲养管理提供一定参考。     

Using video playbacks to study visual communication in a marine fish, Salaria pavo

Video playbacks have been successfully applied to the study of visual communication in several groups of animals. However, this technique is controversial as video monitors are designed with the human visual system in mind. Differences between the visual capabilities of humans and other animals will lead to perceptually different interpretations of video images. We simultaneously presented males and females of the peacock blenny, Salaria pavo, with a live conspecific male and an online video image of the same individual. Video images failed to elicit appropriate responses. Males were aggressive towards the live male but not towards video images of the same male. Similarly, females courted only the live male and spent more time near this stimulus. In contrast, females of the gynogenetic poecilid Poecilia formosa showed an equal preference for a live and video image of a P. mexicana male, suggesting a response to live animals as strong as to video images. We discuss differences between the species that may explain their opposite reaction to video images. Copyright 2000 The Association for the Study of Animal Behaviour.     

The Effects of Social Experience on Aggressive Behavior in the Green Anole Lizard (Anolis carolinensis)

To understand how context-specific aggression emerges from past experience, we examined how consecutive aggressive encounters influence aggressive behavior and stress responses of male green anole lizards ( Anolis carolinensis ). Animals were shown a video clip featuring an aggressively displaying conspecific male, which provoked aggressive responding, while control animals viewed a neutral video. After 5 d of interaction with the videos, both the subject and control groups were presented with a live conspecific. As a non-invasive assay of stress responses, we measured changes in body color and eyespot darkness, two features known to be strongly correlated with titers of stress hormones. Our results demonstrate that experience increased aggression in male anoles, but that increases in aggression to a repeated stimulus were transient. Tests with a novel conspecific indicate that the experienced animals remained aggressive when presented with novel stimuli. Although there were differences in the morphological indicators of the stress response between experimental and control groups during video presentations, there were no differences when presented with novel conspecifics. These data indicate that experience-dependent differences were not mediated by differences in the 'stressfulness' of aggressive interaction, as thought to be the case for animals in chronic subordinate/dominant dyads. We suggest that habituation and reinforcement interact to promote aggressive responding and to restrict it to novel individuals. Such context specificity is a hallmark of natural patterns of aggression in territorial species.     

Individual and social factors affecting the ability of American crows to solve and master a string pulling task

Crows and other birds in the family Corvidae regularly share information to learn the identity and whereabouts of dangerous predators, but can they use social learning to solve a novel task for a food reward? Here, we examined the factors affecting the ability of 27 wild-caught American crows to solve a common string pulling task in a laboratory setting. We split crows into two groups; one group was given the task after repeatedly observing a conspecific model the solution and the other solved in the absence of conspecific models. We recorded the crows’ estimated age, sex, size, body condition, level of nervousness, and brain volume using DICOM images from a CT scan. Although none of these variables were statistically significant, crows without a conspecific model and large brain volumes consistently mastered the task in the minimum number of days, whereas those with conspecific models and smaller brain volumes required varying and sometimes a substantial number of days to master the task. We found indirect evidence that body condition might also be important for motivating crows to solve the task. Crows with conspecific models were no more likely to initially solve the task than those working the puzzle without social information, but those that mastered the task usually copied the method most frequently demonstrated by their knowledgeable neighbors. These findings suggest that brain volume and possibly body condition may be factors in learning new tasks and that crows can use social learning to refine their ability to obtain a novel food source, although they must initially learn to access it themselves.     

Video demonstrations seed alternative problem-solving techniques in wild common marmosets

Studies of social learning and tradition formation under field conditions have recently gained momentum, but suffer from the limited control of socio-ecological factors thought to be responsible for transmission patterns. The use of artificial visual stimuli is a potentially powerful tool to overcome some of these problems. Here, in a field experiment, we used video images of unfamiliar conspecifics performing virtual demonstrations of foraging techniques. We tested 12 family groups of wild common marmosets. Six groups received video demonstrations (footage of conspecifics either pulling a drawer open or pushing a lid upwards, in an ‘artificial fruit’); the other six groups served as controls (exposed to a static image of a conspecific next to the fruit). Subjects in video groups were more manipulative and successful in opening the fruit than controls; they were also more likely to use the technique they had witnessed and thus could serve as live models for other family members. To our knowledge, this is the first study that used video demonstrations in the wild and demonstrated the potent force of social learning, even from unfamiliar conspecifics, under field conditions.     

Effects of visual context upon functional connectivity during observation of biological motions

The aim of this study was to examine brain responses, in particular functional connectivity, to different visual stimuli depicting familiar biological motions. Ten subjects actively observed familiar biological motions embedded in point-light and video displays. Electroencephalograms were recorded from 64 electrodes. Activity was considered in three frequency bands (4-8 Hz, 8-10 Hz, and 10-13 Hz) using a non-linear measure of functional connectivity. In the 4-8 Hz and 8-10 Hz frequency bands, functional connectivity for the SMA was greater during the observation of biological motions presented in a point-light display compared to the observation of motions presented in a video display. The reverse was observed for the 4-8 Hz frequency band for the left temporal area. Explanations related to: (i) the task demands (i.e., attention and mental effort), (ii) the role(s) of theta and alpha oscillations in cognitive processes, and (iii) the function(s) of cortical areas are discussed. It has been suggested that attention was required to process human biological motions under unfamiliar viewing conditions such as point-light display.     

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.     

Spatial and behavioural measures of social discrimination by captive male zebra finches: Implications of sexual and species differences for recognition research

All bird species reproduce sexually and individuals need to correctly identify conspecifics for successful breeding. Captive zebra finches are a model system for studying the factors involved in species recognition and mate choice. However, male zebra finches’ behavioural responses in a spatial preference paradigm to a range of estrildid finch species, other than domesticated Bengalese finches, remain unknown. We investigated spatial and display responses of male zebra finch subjects to stimulus females between conspecific and four phylogeographically relevant finch species, in addition to female Bengalese finches. Surprisingly, male subjects did not show consistent spatial association with conspecific over heterospecific females. Overall, as predicted by sexual selection theory, the spatial proximity responses of males were less discriminatory compared to female zebra finches’ responses tested previously using the same paradigm. However, male subjects showed consistently more behavioural displays towards female conspecifics than heterospecifics which were positively related to the behavioural display rates of the respective female stimuli. Some male behavioural responses, other than song, also showed significant differences between the different stimulus species and consistently differed across individual test subjects, with the most individual subject variation seen in choice trials between female conspecific and Bengalese finch stimuli. The results are important for the design and interpretation of future behavioural and neurobiological experiments on species recognition systems using the zebra finch as a model species.     

Eavesdropping and signal matching in visual courtship displays of spiders

Eavesdropping on communication is widespread among animals, e.g. bystanders observing male-male contests, female mate choice copying and predator detection of prey cues. Some animals also exhibit signal matching, e.g. overlapping of competitors' acoustic signals in aggressive interactions. Fewer studies have examined male eavesdropping on conspecific courtship, although males could increase mating success by attending to others' behaviour and displaying whenever courtship is detected. In this study, we show that field-experienced male Schizocosa ocreata wolf spiders exhibit eavesdropping and signal matching when exposed to video playback of courting male conspecifics. Male spiders had longer bouts of interaction with a courting male stimulus, and more bouts of courtship signalling during and after the presence of a male on the video screen. Rates of courtship (leg tapping) displayed by individual focal males were correlated with the rates of the video exemplar to which they were exposed. These findings suggest male wolf spiders might gain information by eavesdropping on conspecific courtship and adjust performance to match that of rivals. This represents a novel finding, as these behaviours have previously been seen primarily among vertebrates.     

Why do chimpanzees have diverse behavioral repertoires yet lack more complex cultures? Invention and social information use in a cumulative task

Humans are distinctive in their dependence upon products of culture for survival, products that have evolved cumulatively over generations such that many cannot now be created by a single individual. Why the cultural capacity of humans appears unrivalled in the animal kingdom is a topic of ongoing debate. Here we explore whether innovation and/or social learning propensities may constrain the ability of one of our closest living relatives, chimpanzees (Pan troglodytes), to master an extractive foraging and tool-use task designed to afford opportunities for cumulative culture to develop. We further explore the potential demographic characteristics associated with novel task solutions. Chimpanzees (N = 53) were inventive, flexibly exploring the novel task, albeit complex inventions were rare and shaped by prior individual experience with similar tool-use tasks. However, they displayed no evidence of cumulative cultural learning. Communities displayed richer behavioral repertoires and had greater task success than chimpanzees tested in an asocial control condition, but their solution complexity did not surpass what individuals invented. The lack of social transmission of complex and beneficial solutions in contexts like those we studied provides one explanation for the limited cumulative culture observed in this species.