Finger drawing by infant chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>)

We introduced a new technique to investigate the development of scribbling in very young infants. We tested three infant chimpanzees to compare the developmental processes of scribbling between humans and chimpanzees. While human infants start to scribble on paper at around the age of 18 months, our 13- to 23-month-old infant chimpanzees had never been observed scribbling prior to this study. We used a notebook computer with a touch-sensitive screen. This apparatus was able to record the location of the subjects touches on the screen. Each touch generated a fingertip-sized dot at the corresponding on-screen location. During spontaneous interactions with this apparatus, all three infants and two mother chimpanzees left scribbles with their fingers on the screen. The scribbles contained not only simple dots or short lines, but also curves and hook-like lines or loops, most of which were observed in the instrumental drawings of adult chimpanzees. The results suggest that perceptual-motor control for finger drawing develops in infant chimpanzees. Two of the infants performed their first scribble with a marker on paper at the age of 20–23 months. Just prior to this, they showed a rapid increase in combinatory manipulation of objects. These findings suggest that the development of combinatory manipulation of objects as well as that of perceptual-motor control may be necessary for the emergence of instrumental drawing on paper.     

Recognition of pictorial representations by chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>)

In this study, I investigated chimpanzees' ability to recognize pictorial representations. Four adults and three juvenile chimpanzees were trained to choose images of photographs of flowers among 12 items belonging to four categories on a touch-sensitive monitor. As a generalization test, the following five types of images were presented: (1) novel photographs, (2) colored sketches (more realistic), (3) a colored clip art (cartoon-like images), (4) black-and-white line drawings, and (5) Kanji characters (as the control images). One adult and all three juvenile chimpanzees were able to choose any style of the nonphotographic images of flowers significantly above the chance level, whereas none could choose the correct Kanji characters corresponding to a flower significantly above the chance level. The other three adult chimpanzees' performance level did not exceed the chance level in terms of choosing nonphotographic images although they showed good transfer skills to novel photographs. The results revealed that not all chimpanzees could recognize pictures used by humans without training. The results also suggest "critical period" in acquisition of skill in recognizing pictures in chimpanzees. Only one adult chimpanzee, who had acquired skill in recognizing visual symbols, also recognized pictures aside from the juvenile chimpanzees. Her learning history might have aided her in acquiring this skill. The results of this study suggest a relationship between pictorial competence and symbolic one.     

Subgenual cingulate cortex and personality in chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>)

Animals vary in their dispositions, abilities, and moods and demonstrate characteristic behavior patterns that remain consistent across situation and time. This study describes the relationship between measures of personality in the chimpanzee and the structure of the subgenual cingulate cortex (SGCC). Measures of individual traits and personality factors (dominance, extraversion, conscientiousness, and agreeableness) and assessments of percentage of SGCC gray matter (PGM) and asymmetry taken from MRI scans were obtained for 74 chimpanzees housed at the Yerkes National Primate Research Center. PGM in the SGCC was significantly higher for females than for males and was significantly correlated with two personality factors (dominance and conscientiousness) in male apes. There was also a population-level leftward asymmetry in the SGCC. These results are discussed in terms of current models of SGCC function, which suggest that this area may play a role in the biological foundation of personality.     

Discrimination of contour-deleted images in baboons (<Emphasis Type="Italic">Papio papio</Emphasis>) and chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>)

Humans readily group component elements into a coherent perceptual whole and perceive the global form of visual patterns in priority over local features, which stands in contrast to at least some data from the animal literature, suggesting possible species differences in perceptual processes. In this study, chimpanzees and baboons were required to match intact and partially contour-deleted line-drawings in a computerized task in order to further explore the ability of nonhuman primates to group component elements into a coherent perceptual whole and to determine to what extent they use the global form or the local features. Experiment 1 showed that the baboons and chimpanzees matched intact continuous line-drawings (intact-intact) more easily than partially contour-deleted line-drawings (deleted-deleted). Both species could also match partially deleted line-drawings with their intact version (deleted-intact), but at a lower performance level. Experiment 2 further showed that subjects from the two species could match partially deleted coherent line-drawings (deleted-deleted) more easily than their scrambled incoherent versions (scrambled-scrambled). They could however match the coherent deleted forms with their scrambled version (deleted-scrambled). It is suggested that solutions of these tasks rely on the processing of both global and local cues, with no clear-cut species difference in that ability. Overall, the results show that contour completion of line-drawings was not easy. Implications on the processing of human-made two-dimensional representations such as line-drawings are discussed.     

Perception of depth from shading in infant chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>)

We investigated the ability to perceive depth from shading, one of the pictorial depth cues, in three chimpanzee infants aged 4–10 months old, using a preferential reaching task commonly used to study pictorial depth perception in human infants. The chimpanzee infants reached significantly more to three-dimensional toys than to pictures thereof and more to the three-dimensional convex than to the concave. Furthermore, two of the three infants reached significantly more to the photographic convex than to the photographic concave. These infants also looked longer at the photographic convex than the concave. Our results suggest that chimpanzees perceive, at least as early as the latter half of the first year of life, pictorial depth defined by shading information. Photographic convexes contain richer information about pictorial depth (e.g., attached shadow, cast shadow, highlighted area, and global difference in brightness) than simple computer-graphic graded patterns. These cues together might facilitate the infants perception of depth from shading.     

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.     

Geometric distortions affect face recognition in chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>) and monkeys (<Emphasis Type="Italic">Macaca mulatta</Emphasis>)

All primates can recognize faces and do so by analyzing the subtle variation that exists between faces. Through a series of three experiments, we attempted to clarify the nature of second-order information processing in nonhuman primates. Experiment one showed that both chimpanzees (Pan troglodytes) and rhesus monkeys (Macaca mulatta) tolerate geometric distortions along the vertical axis, suggesting that information about absolute position of features does not contribute to accurate face recognition. Chimpanzees differed from monkeys, however, in that they were more sensitive to distortions along the horizontal axis, suggesting that when building a global representation of facial identity, horizontal relations between features are more diagnostic of identity than vertical relations. Two further experiments were performed to determine whether the monkeys were simply less sensitive to horizontal relations compared to chimpanzees or were instead relying on local features. The results of these experiments confirm that monkeys can utilize a holistic strategy when discriminating between faces regardless of familiarity. In contrast, our data show that chimpanzees, like humans, use a combination of holistic and local features when the faces are unfamiliar, but primarily holistic information when the faces become familiar. We argue that our comparative approach to the study of face recognition reveals the impact that individual experience and social organization has on visual cognition.     

Robust representations of individual faces in chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>) but not monkeys (<Emphasis Type="Italic">Macaca mulatta</Emphasis>)

Being able to recognize the faces of our friends and family members no matter where we see them represents a substantial challenge for the visual system because the retinal image of a face can be degraded by both changes in the person (age, expression, pose, hairstyle, etc.) and changes in the viewing conditions (direction and degree of illumination). Yet most of us are able to recognize familiar people effortlessly. A popular theory for how face recognition is achieved has argued that the brain stabilizes facial appearance by building average representations that enhance diagnostic features that reliably vary between people while diluting features that vary between instances of the same person. This explains why people find it easier to recognize average images of people, created by averaging multiple images of the same person together, than single instances (i.e. photographs). Although this theory is gathering momentum in the psychological and computer sciences, there is no evidence of whether this mechanism represents a unique specialization for individual recognition in humans. Here we tested two species, chimpanzees (Pan troglodytes) and rhesus monkeys (Macaca mulatta), to determine whether average images of different familiar individuals were easier to discriminate than photographs of familiar individuals. Using a two-alternative forced-choice, match-to-sample procedure, we report a behaviour response profile that suggests chimpanzees encode the faces of conspecifics differently than rhesus monkeys and in a manner similar to humans.     

How chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>) perform in a modified emotional Stroop task

The emotional Stroop task is an experimental paradigm developed to study the relationship between emotion and cognition. Human participants required to identify the color of words typically respond more slowly to negative than to neutral words (emotional Stroop effect). Here we investigated whether chimpanzees (Pan troglodytes) would show a comparable effect. Using a touch screen, eight chimpanzees were trained to choose between two simultaneously presented stimuli based on color (two identical images with differently colored frames). In Experiment 1, the images within the color frames were shapes that were either of the same color as the surrounding frame or of the alternative color. Subjects made fewer errors and responded faster when shapes were of the same color as the frame surrounding them than when they were not, evidencing that embedded images affected target selection. Experiment 2, a modified version of the emotional Stroop task, presented subjects with four different categories of novel images: three categories of pictures of humans (veterinarian, caretaker, and stranger), and control stimuli showing a white square. Because visits by the veterinarian that include anaesthetization can be stressful for subjects, we expected impaired performance in trials presenting images of the veterinarian. For the first session, we found correct responses to be indeed slower in trials of this category. This effect was more pronounced for subjects whose last anaesthetization experience was more recent, indicating that emotional valence caused the slowdown. We propose our modified emotional Stroop task as a simple method to explore which emotional stimuli affect cognitive performance in nonhuman primates.     

Long-term retention of the differential values of Arabic numerals by chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>)

As previously reported (Beran and Rumbaugh, 2001), two chimpanzees used a joystick to collect dots, one-at-a-time, on a computer monitor, and then ended a trial when the number of dots collected was equal to the Arabic numeral presented for the trial. Here, the chimpanzees were presented with the task again after an interval of 6 months and then again after an additional interval of 3.25 years. During each interval, the chimpanzees were not presented with the task, and this allowed an assessment of the extent to which both animals retained the values of each Arabic numeral. Despite lower performance at each retention interval compared to the original study, both chimpanzees performed above chance levels in collecting a quantity of dots equal to the target numeral, one chimpanzee for the numerals 1-7, and the second chimpanzee for the numerals 1-6. For the 3.25-year retention, errors were more dispersed around each target numeral than in the original study, but the chimpanzees' performances again appeared to be based on a continuous representation of magnitude rather than a discrete representation of number. These data provide an experimental demonstration of long-term retention of the differential values of Arabic numerals by chimpanzees.     

Development of interception of moving targets by chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>) in an automated task

The experiments investigated how two adult captive chimpanzees learned to navigate in an automated interception task. They had to capture a visual target that moved predictably on a touch monitor. The aim of the study was to determine the learning stages that led to an efficient strategy of intercepting the target. The chimpanzees had prior training in moving a finger on a touch monitor and were exposed to the interception task without any explicit training. With a finger the subject could move a small "ball" at any speed on the screen toward a visual target that moved at a fixed speed either back and forth in a linear path or around the edge of the screen in a rectangular pattern. Initial ball and target locations varied from trial to trial. The subjects received a small fruit reinforcement when they hit the target with the ball. The speed of target movement was increased across training stages up to 38 cm/s. Learning progressed from merely chasing the target to intercepting the target by moving the ball to a point on the screen that coincided with arrival of the target at that point. Performance improvement consisted of reduction in redundancy of the movement path and reduction in the time to target interception. Analysis of the finger's movement path showed that the subjects anticipated the target's movement even before it began to move. Thus, the subjects learned to use the target's initial resting location at trial onset as a predictive signal for where the target would later be when it began moving. During probe trials, where the target unpredictably remained stationary throughout the trial, the subjects first moved the ball in anticipation of expected target movement and then corrected the movement to steer the ball to the resting target. Anticipatory ball movement in probe trials with novel ball and target locations (tested for one subject) showed generalized interception beyond the trained ball and target locations. The experiments illustrate in a laboratory setting the development of a highly complex and adaptive motor performance that resembles navigational skills seen in natural settings where predators intercept the path of moving prey. Electronic Supplementary Material Supplementary material is available for this article if you access the article at . A link in the frame on the left on that page takes you directly to the supplementary material.     

Matching vocalizations to vocalizing faces in a chimpanzee (<Emphasis Type="Italic">Pan troglodytes</Emphasis>)

Auditory–visual processing of species-specific vocalizations was investigated in a female chimpanzee named Pan. The basic task was auditory–visual matching-to-sample, where Pan was required to choose the vocalizer from two test movies in response to a chimpanzees vocalization. In experiment 1, movies of vocalizing and silent faces were paired as the test movies. The results revealed that Pan recognized the status of other chimpanzees whether they vocalized or not. In experiment 2, two different types of vocalizing faces of an identical individual were prepared as the test movies. Pan recognized the correspondence between vocalization types and faces. These results suggested that chimpanzees possess crossmodal representations of their vocalizations, as do humans. Together with the ability of vocal individual recognition, this ability might reflect chimpanzees profound understanding of the status of other individuals.     

Enhanced recognition of emotional stimuli in the chimpanzee (<Emphasis Type="Italic">Pan troglodytes</Emphasis>)

Humans often have a better memory of emotional events than neutral events. From the comparative-cognitive perspective, we explored the enhancement of recognition memory by emotion in chimpanzees (Pan troglodytes) using a serial probe recognition task. In this task, we sequentially presented a list of pictures to subjects and then tested their recognition of specific pictures from within the list. We selected pictures of aggressive chimpanzees as emotional stimuli and less tensed, relaxed chimpanzees as neutral stimuli. In Experiment 1, we gave four-item lists to two young chimpanzees; one showed significantly greater recognition of pictures depicting aggressive chimpanzees than neutral ones. In Experiment 2, this chimpanzee was further tested using a recognition task with eight-item lists. The subject again showed better recognition of emotional stimuli than neutral. Furthermore, the presence of an emotional stimulus in the list also facilitated recognition of the neutral item immediately following it. Overall, although only one of the two chimpanzees showed enhanced recognition memory by emotional stimuli, this is the first demonstration of such a response in the chimpanzee. The findings are discussed in comparison with those of human studies.     

Perceptual biases for multimodal cues in chimpanzee (<Emphasis Type="Italic">Pan troglodytes</Emphasis>) affect recognition

The ability of organisms to discriminate social signals, such as affective displays, using different sensory modalities is important for social communication. However, a major problem for understanding the evolution and integration of multimodal signals is determining how humans and animals attend to different sensory modalities, and these different modalities contribute to the perception and categorization of social signals. Using a matching-to-sample procedure, chimpanzees discriminated videos of conspecifics' facial expressions that contained only auditory or only visual cues by selecting one of two facial expression photographs that matched the expression category represented by the sample. Other videos were edited to contain incongruent sensory cues, i.e., visual features of one expression but auditory features of another. In these cases, subjects were free to select the expression that matched either the auditory or visual modality, whichever was more salient for that expression type. Results showed that chimpanzees were able to discriminate facial expressions using only auditory or visual cues, and when these modalities were mixed. However, in these latter trials, depending on the expression category, clear preferences for either the visual or auditory modality emerged. Pant-hoots and play faces were discriminated preferentially using the auditory modality, while screams were discriminated preferentially using the visual modality. Therefore, depending on the type of expressive display, the auditory and visual modalities were differentially salient in ways that appear consistent with the ethological importance of that display's social function.     

Chimpanzee (<Emphasis Type="Italic">Pan troglodytes</Emphasis>) and orangutan (<Emphasis Type="Italic">Pongo abelii</Emphasis>) forethought: self-control and pre-experience in the face of future tool use

Planning for future needs has traditionally been considered to be restricted to human cognition. Although recent studies on great ape and corvid cognition challenge this belief, the phylogenesis of human planning remains largely unknown. The complex skill for future planning has not yet been satisfactorily established in any other extant primate species than our own. In humans, planning for future needs rely heavily on two overarching capacities, both of which lie at the heart of our cognition: self-control, often defined as the suppression of immediate drives in favor of delayed rewards, and mental time travel, which could be described as a detached mental experience of a past or future event. Future planning is linked to additional high complexity cognition such as metacognition and a consciousness usually not attributed to animals. In a series of four experiments based on tool use, we demonstrate that chimpanzees (Pan troglodytes) and orangutans (Pongo abelii) override immediate drives in favor of future needs, and they do not merely rely on associative learning or semantic prospection when confronted with a planning task. These results suggest that great apes engage in planning for the future by out competing current drives and mentally pre-experiencing an upcoming event. This suggests that the advanced mental capacities utilized in human future planning are shared by phylogenetically more ancient species than previously believed.     

Combinations of <Emphasis Type="Italic">Stit</Emphasis> with <Emphasis Type="Italic">Ought</Emphasis> and <Emphasis Type="Italic">Know</Emphasis>

This paper presents a short survey of recent developments in stit theories, with an emphasis on combinations of stit and deontic logic, and those of stit and epistemic logic.     

Chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>) show the isolation effect during serial list recognition memory tests

The isolation effect (or von Restorff effect) occurs when one item in a to-be-remembered list is distinctive from all remaining items, and memory for that item is enhanced. Four chimpanzees were presented with a serial list of four photographs. In the homogeneous condition, all list items were from the same semantic category (e.g., four fruits). In the isolate condition, three items were from the same category, but the fourth item (the isolate) was from a different category (e.g., three fruits and one toy). Then, two photographs were presented, and the chimpanzees had to select the one that was from the list. Two of four chimpanzees were significantly more likely to select a correct isolate item than an item from the same list position in the homogeneous condition for at least some list positions. This facilitation in performance was for isolate items only, as presenting an isolate item in a list did not facilitate greater recognition of other list items compared to the homogeneous condition. These results indicated that some chimpanzees perceived the semantic categories of the photographs, and categorization of photographs led to the isolation effect. Thus, chimpanzees may share with humans some aspects of memory organization that involve spontaneously categorizing visual stimuli and recognizing categorically unique stimuli.     

<Emphasis Type="Italic">Ziran</Emphasis> and <Emphasis Type="Italic">Wuwei</Emphasis> in the <Emphasis Type="Italic">Daodejing</Emphasis>: An Ethical Assessment

In Daoist philosophy, the self is understood as an individual interdependent with others, and situated within a broader environment. Within this framework, the concept ziran is frequently understood in terms of naturalness or nature while wuwei is explained in terms of non-oppressive government. In many existing accounts, little is done to connect these two key Daoist concepts. Here, I suggest that wuwei and ziran are correlated, ethical, concepts. Together, they provide a unifying ethical framework for understanding the philosophy of the Daodejing. I explore the meaning of ziran as self-so-ness or, in human terms, as pertaining to an individual’s spontaneity. The appropriate response to the spontaneity of individuals is to avoid, insofar as possible, imposing or using restrictive norms and methods, that is, wuwei. According to this view, ziran and wuwei offer an account of ethics that attends to core notions of interdependent selfhood, including mutuality, relationality, interdependence, symbiosis, and responsiveness.     

Comprehension of human pointing gestures in young human-reared wolves (<Emphasis Type="Italic">Canis lupus</Emphasis>) and dogs (<Emphasis Type="Italic">Canis familiaris</Emphasis>)

Dogs have a remarkable skill to use human-given cues in object-choice tasks, but little is known to what extent their closest wild-living relative, the wolf can achieve this performance. In Study 1, we compared wolf and dog pups hand-reared individually and pet dogs of the same age in their readiness to form eye-contact with a human experimenter in an object-choice task and to follow her pointing gesture. The results showed that dogs already at 4 months of age use momentary distal pointing to find hidden food even without intensive early socialization. Wolf pups, on the contrary, do not attend to this subtle pointing. Accordingly in Studies 2 and 3, these wolves were tested longitudinally with this and four other (easier) human-given cues. This revealed that wolves socialized at a comparable level to dogs are able to use simple human-given cues spontaneously if the human’s hand is close to the baited container (e.g. touching, proximal pointing). Study 4 showed that wolves can follow also momentary distal pointing similarly to dogs if they have received extensive formal training. Comparing the wolves to naïve pet dogs of the same age revealed that during several months of formal training wolves can reach the level of dogs in their success of following momentary distal pointing in parallel with improving their readiness to form eye-contact with a human experimenter. We assume that the high variability in the wolves’ communicative behaviour might have provided a basis for selection during the course of domestication of the dog.