Assessing the motivational value of infant video clips on chimpanzees through discrimination learning task.

The motivational value of visual infant stimuli in humans is considered to encourage parental behavior. To explore the evolutionary roots of this preference for infants, we examined the reward value of conspecific infant videos compared to adult ones in nine chimpanzees. We employed a novel approach, a simultaneous discrimination task with differential sensory reinforcement. In Experiments 1 and 2, we tested if watching conspecific infant videos is more rewarding than watching adult ones. Participants were required to discriminate between two visual stimuli by a touch panel task. In video reward trials, a video clip featuring a chimpanzee infant followed a correct choice, while one featuring an adult followed an incorrect choice. However, the percentage of correct choices did not significantly differ from chance except in one chimpanzee, indicating that chimpanzees did not exhibit a preference for watching infant videos over those of adult. In Experiment 3, we tested if chimpanzees prefer conspecific videos over a blank screen; however, we did not find evidence either at a group level. These results suggest that the incentive salience of infant stimuli may not be universally compelling across species. Additionally, we discuss the limitations of the task using sensory reinforcement.

Do chimpanzees see a face on Mars? A search for face pareidolia in chimpanzees.

We sometimes perceive meaningful patterns or images in random arrangements of colors and shapes. This phenomenon is called pareidolia and has recently been studied intensively, especially face pareidolia. In contrast, there are few comparative-cognitive studies on face pareidolia with nonhuman primates. This study explored behavioral evidence for face pareidolia in chimpanzees using visual search and matching tasks. Faces are processed in a configural manner, and their perception and recognition are hampered by inversion and misalignment of top and bottom parts. We investigated whether the same effect occurs in a visual search for face-like objects. The results showed an effect of misalignment. On the other hand, consistent results were not obtained with the photographs of fruits. When only the top or bottom half of the face-like object was presented, chimpanzees showed better performance for the top-half condition, suggesting the importance of the eye area in face pareidolia. In the positive-control experiments, chimpanzees received the same experiment using human faces and human participants with face-like objects and fruits. As a result, chimpanzees showed an inefficient search for inverted and misaligned faces and humans for manipulated face-like objects. Finally, to examine the role of face awareness, we tested matching a human face to a face-like object in chimpanzees but obtained no substantial evidence that they saw the face-like object as a "face." Based on these results, we discussed the extents and limits of face pareidolia in chimpanzees.

Slowly walking down to the more food: relative quantity discrimination in African spurred tortoises (Centrochelys sulcata).

Quantity discrimination, is thought to be highly adaptive as it allows an organism to select greater amounts of food or larger social groups. In contrast to mammals, the processes underlying this ability are not as well understood in reptiles. This study examined the effects of ratio and number size on relative quantity discrimination in African spurred tortoises (Centrochelys sulcata). To assess these effects, tortoises were presented with trays containing favored food pieces in all possible number combinations between 1 and 7. The tortoises had to approach the tray they perceived as having the larger quantity. If correct, they received one piece of food as reinforcement. The results revealed that relative quantity discrimination was influenced by the ratio between the numbers of pieces, with performance improving as the ratio between the numbers increased. This finding suggests that the approximate number system or analogue magnitude estimation may control their behavior. However, as the number size increased, their performance declined, also suggesting that the approximate number system alone could not explain the present results.

Left or right, that is the question: use of egocentric frame of reference and the right-eye advantage for understanding gestural signs in bottlenose dolphins (Tursiops truncatus).

How do bottlenose dolphins visually perceive the space around them? In particular, what cues do they use as a frame of reference for left-right perception? To address this question, we examined the dolphin's responses to various manipulations of the spatial relationship between the dolphin and the trainer by using gestural signs for actions given by the trainer, which have different meanings in the left and right hands. When the dolphins were tested with their backs to the trainer (Experiment 1) or in an inverted position underwater (Experiments 2 and 3), correct responses from the trainer's perspective were maintained for signs related to movement direction instructions. In contrast, reversed responses were frequently observed for signs that required different sounds for the left and right hands. When the movement direction instructions were presented with symmetrical graphic signs such as " × " and "●", accuracy decreased in the inverted posture (Experiment 3). Furthermore, when the signs for sounds were presented from either the left or right side of the dolphin's body, performance was better when the side of the sign movement coincided with the body side on which it was presented than when it was mismatched (Experiment 4). In the final experiment, when one eye was covered with an eyecup, the results showed that, as in the case of body-side presentation, performance was better when the open eye coincided with the side on which the sign movement was presented. These results indicate that dolphins used the egocentric frame for visuospatial cognition. In addition, they showed better performances when the gestural signs were presented to the right eye, suggesting the possibility of a left-hemispheric advantage in the dolphin's visuospatial cognition.

Don't look back on failure: spontaneous uncertainty monitoring in chimpanzees.

During computer-controlled cognitive tasks, chimpanzees often look up at the food dispenser, which activates at the same time as feedback for the correct choice but not for feedback for the incorrect choice. Do these "looking back" behaviors also indicate signs of spontaneous monitoring of their confidence in their choices? To address this question, we delayed the feedback for 1 s after their choice responses and observed their look-back behaviors during the delay period. Two chimpanzees looked up at the food dispenser significantly less frequently when their choice was incorrect (but the feedback was not given) than when it was correct. These look-back behaviors have not been explicitly trained under experimental contexts. Therefore, these results indicate that chimpanzees spontaneously change the frequency of their look-back behaviors in response to the correctness or incorrectness of their own choices, even without external feedback, suggesting that their look-back behaviors may reflect the level of "confidence" or "uncertainty" of their responses immediately before.

Attention to inattention: Effect of the other's attentional state on visual search performance in chimpanzees.

In the present study, we examined the effects of the other's triadic attention to objects on visual search performances in chimpanzees. We found the search-asymmetry-like effect of the other's attentional state; the chimpanzees searched a target object not attended by the other individual more efficiently than that attended (Experiment 1). Additional experiments explored the possibility that the other individual "holding an object but not looking at it" led to expectancy violation (Experiment 2) or the role of nonsocial cues such as the proximity relation between the head and the object (Experiment 3). Still, these accounts alone did not explain this effect. It was also shown that the other's attentional state affected the chimpanzees' performances more readily as the interference effect than the facilitation effect (Experiment 4). Furthermore, the same effect was observed in the visual search for the gaze (head direction) of others (Experiment 5). We obtained the same results using photographs of chimpanzees (Experiment 6). Contrary to the chimpanzees, humans detected the object to which attention was directed more efficiently than vice versa (Experiment 7). The present results may reflect species differences between chimpanzees and humans in processing triadic social attention.

Rhythmic swaying induced by sound in chimpanzees (Pan troglodytes).

Music and dance are universal across human culture and have an ancient history. One characteristic of music is its strong influence on movement. For example, an auditory beat induces rhythmic movement with positive emotions in humans from early developmental stages. In this study, we investigated if sound induced spontaneous rhythmic movement in chimpanzees. Three experiments showed that: 1) an auditory beat induced rhythmic swaying and other rhythmic movements, with larger responses from male chimpanzees than female chimpanzees; 2) random beat as well as regular beat induced rhythmic swaying and beat tempo affected movement periodicity in a chimpanzee in a bipedal posture; and 3) a chimpanzee showed close proximity to the sound source while hearing auditory stimuli. The finding that male chimpanzees showed a larger response to sound than female chimpanzees was consistent with previous literature about "rain dances" in the wild, where male chimpanzees engage in rhythmic displays when hearing the sound of rain starting. The fact that rhythmic swaying was induced regardless of beat regularity may be a critical difference from humans, and a further study should reveal the physiological properties of sound that induce rhythmic movements in chimpanzees. These results suggest some biological foundation for dancing existed in the common ancestor of humans and chimpanzees ∼6 million years ago. As such, this study supports the evolutionary origins of musicality.

Chimpanzees (Pan troglodytes) detect strange body parts: an eye-tracking study.

This study investigated chimpanzee body representation by testing whether chimpanzees detect strangeness in body parts. We tested six chimpanzees with edited chimpanzee body pictures in eye-tracking tasks. The target body parts were arms or legs. For either target, there were four conditions: "normal" condition as control, where all bodies were normal; "misplaced" condition, where one arm or one leg was misplaced to an incorrect body location in each picture; "replaced by a chimpanzee part" condition, where one arm or one leg was replaced by a chimpanzee leg or arm, respectively, in its original place in each picture; and "replaced by a human part" condition, where one arm or one leg was replaced by a human arm or leg in each picture. Compared to the looking times toward the normal parts, chimpanzees had significantly longer looking times toward the human arms or legs. The looking times toward the misplaced parts were also longer than the normal parts, but the difference just failed to meet significance. These results indicate more interests toward strange body parts, compared to typical parts, suggesting that chimpanzees might have a body representation that is sufficiently sensitive to detect these aspects of strangeness.

No evidence of spatial representation of age, but "own-age bias" like face processing found in chimpanzees.

Previous studies have revealed that non-human primates can differentiate the age category of faces. However, the knowledge about age recognition in non-human primates is very limited and whether non-human primates can process facial age information in a similar way to humans is unknown. As humans have an association between time and space (e.g., a person in an earlier life stage to the left and a person in a later life stage to the right), we investigated whether chimpanzees spatially represent conspecifics' adult and infant faces. Chimpanzees were tested using an identical matching-to-sample task with conspecific adult and infant face stimuli. Two comparison images were presented vertically (Experiment 1) or horizontally (Experiment 2). We analyzed whether the response time was influenced by the position and age category of the target stimuli, but there was no evidence of correspondence between space and adult/infant faces. Thus, evidence of the spatial representation of the age category was not found. However, we did find that the response time was consistently faster when they discriminated between adult faces than when they discriminated between infant faces in both experiments. This result is in line with a series of human face studies that suggest the existence of an "own-age bias." As far as we know, this is the first report of asymmetric face processing efficiency between infant and adult faces in non-human primates.

Great apes use self-experience to anticipate an agent's action in a false-belief test.

Human social life depends on theory of mind, the ability to attribute mental states to oneself and others. A signature of theory of mind, false belief understanding, requires representing others' views of the world, even when they conflict with one's own. After decades of research, it remains controversial whether any nonhuman species possess a theory of mind. One challenge to positive evidence of animal theory of mind, the behavior-rule account, holds that animals solve such tasks by responding to others' behavioral cues rather than their mental states. We distinguish these hypotheses by implementing a version of the "goggles" test, which asks whether, in the absence of any additional behavioral cues, animals can use their own self-experience of a novel barrier being translucent or opaque to determine whether another agent can see through the same barrier. We incorporated this paradigm into an established anticipatory-looking false-belief test for great apes. In a between-subjects design, apes experienced a novel barrier as either translucent or opaque, although both looked identical from afar. While being eye tracked, all apes then watched a video in which an actor saw an object hidden under 1 of 2 identical boxes. The actor then scuttled behind the novel barrier, at which point the object was relocated and then removed. Only apes who experienced the barrier as opaque visually anticipated that the actor would mistakenly search for the object in its previous location. Great apes, therefore, appeared to attribute differential visual access based specifically on their own past perceptual experience to anticipate an agent's actions in a false-belief test.

How chimpanzees and children perceive other species' bodies: Comparing the expert effect.

Human adults are better at recognizing upright bodies than inverted bodies. This inversion effect is also found for objects with which they have expertise, which is called the expert effect. This study aims to investigate its evolutionary and developmental aspects by testing humans' closest relatives, chimpanzees, and preschool children. Chimpanzees show the inversion effect to chimpanzee bodies, but it is not clear how they perceive other species' bodies. We tested seven chimpanzees in matching-to-sample tasks on touch screens using upright and inverted stimuli, and examined their accuracy and response time. In a previous study, they did not show the inversion effect for bipedal humans in unfamiliar postures, but here in this study they showed it to bipedal humans with familiar postures or crawling postures. This suggests the existence of the expert effect in non-human primates, and that visual or embodied experience is needed to invoke it. It is also supported by the inversion effect they exhibit for horses who share quadrupedal postures, but which they have never seen. Additionally, for conspecifics, the inversion effect was shown regardless of the postures. We tested 33 preschool children using a similar method. They showed the inversion effect to human bodies, but not houses, suggesting the configural processing for bodies, which is found to be stable at the preschool stage. They also showed the inversion effect for chimpanzees and horses, indicating the important role of experience in shaping the ways of object processing.

Underestimating Kanzi? Exploring Kanzi-Oldowan comparisons in light of recent human stone tool replication.

The knapping experiments with Kanzi, a bonobo, are among the most insightful experiments into Oldowan technology ever undertaken. Comparison of his artifacts against archeological material, however, indicated he did not produce Oldowan lithic attributes precisely, prompting suggestions that this indicated cognitive or biomechanical impediments. The literature describing the learning environment provided to Kanzi, we suggest, indicates alternative factors. Based on consideration of wild chimpanzee learning environments, and experiments with modern knappers that have looked at learning environment, we contend that Kanzi's performance was impeded by an impoverished learning environment compared to those experienced by novice Oldowan knappers. Such issues are precisely those that might be tested via a repeat study, but in this case, practical and ethical constraints likely impede this possibility. We propose experiments that may be relevant to drawing conclusions from Kanzi's experiments that may not need to use non-human primates, thus bypassing some of these issues.

The gesture 'Touch': Does meaning-making develop in chimpanzees' use of a very flexible gesture?

In this bottom-up study of gesture, we focused on the details of a single gesture, Touch. We compared characteristics of use by three young chimpanzees with those of 11 adults, their interactive partners, housed in a semi-natural social group at the Kyoto University Primate Research Institute (KUPRI) in Japan. Five hundred eighty-one observations of the gesture Touch were collected across a four-year time span. This single gesture had 36 different forms, was directed to 70 different target locations on the body of social partners, and occurred in 26 different contexts. Significant differences were found between infant and adult initiators in the form, target locations, and contexts of the gesture Touch. There was a wide diversity in form-location patterns within each context, and there were no form-location patterns specific to particular contexts. Thus, we demonstrate that this gesture exhibits flexibility in form and flexibility in use. The results from this study illustrate the importance of contextualized meaning in understanding flexibility in the gesture use of great apes.

Human ostensive signals do not enhance gaze following in chimpanzees, but do enhance object-oriented attention.

The previous studies have shown that human infants and domestic dogs follow the gaze of a human agent only when the agent has addressed them ostensively-e.g., by making eye contact, or calling their name. This evidence is interpreted as showing that they expect ostensive signals to precede referential information. The present study tested chimpanzees, one of the closest relatives to humans, in a series of eye-tracking experiments using an experimental design adapted from these previous studies. In the ostension conditions, a human actor made eye contact, called the participant's name, and then looked at one of two objects. In the control conditions, a salient cue, which differed in each experiment (a colorful object, the actor's nodding, or an eating action), attracted participants' attention to the actor's face, and then the actor looked at the object. Overall, chimpanzees followed the actor's gaze to the cued object in both ostension and control conditions, and the ostensive signals did not enhance gaze following more than the control attention-getters. However, the ostensive signals enhanced subsequent attention to both target and distractor objects (but not to the actor's face) more strongly than the control attention-getters-especially in the chimpanzees who had a close relationship with human caregivers. We interpret this as showing that chimpanzees have a simple form of communicative expectations on the basis of ostensive signals, but unlike human infants and dogs, they do not subsequently use the experimenter's gaze to infer the intended referent. These results may reflect a limitation of non-domesticated species for interpreting humans' ostensive signals in inter-species communication.

Perception of the average size of multiple objects in chimpanzees (Pan troglodytes).

Humans can extract statistical information, such as the average size of a group of objects or the general emotion of faces in a crowd without paying attention to any individual object or face. To determine whether summary perception is unique to humans, we investigated the evolutional origins of this ability by assessing whether chimpanzees, which are closely related to humans, can also determine the average size of multiple visual objects. Five chimpanzees and 18 humans were able to choose the array in which the average size was larger, when presented with a pair of arrays, each containing 12 circles of different or the same sizes. Furthermore, both species were more accurate in judging the average size of arrays consisting of 12 circles of different or the same sizes than they were in judging the average size of arrays consisting of a single circle. Our findings could not be explained by the use of a strategy in which the chimpanzee detected the largest or smallest circle among those in the array. Our study provides the first evidence that chimpanzees can perceive the average size of multiple visual objects. This indicates that the ability to compute the statistical properties of a complex visual scene is not unique to humans, but is shared between both species.

Impaired Air Conditioning within the Nasal Cavity in Flat-Faced Homo.

We are flat-faced hominins with an external nose that protrudes from the face. This feature was derived in the genus Homo, along with facial flattening and reorientation to form a high nasal cavity. The nasal passage conditions the inhaled air in terms of temperature and humidity to match the conditions required in the lung, and its anatomical variation is believed to be evolutionarily sensitive to the ambient atmospheric conditions of a given habitat. In this study, we used computational fluid dynamics (CFD) with three-dimensional topology models of the nasal passage under the same simulation conditions, to investigate air-conditioning performance in humans, chimpanzees, and macaques. The CFD simulation showed a horizontal straight flow of inhaled air in chimpanzees and macaques, contrasting with the upward and curved flow in humans. The inhaled air is conditioned poorly in humans compared with nonhuman primates. Virtual modifications to the human external nose topology, in which the nasal vestibule and valve are modified to resemble those of chimpanzees, change the airflow to be horizontal, but have little influence on the air-conditioning performance in humans. These findings suggest that morphological variation of the nasal passage topology was only weakly sensitive to the ambient atmosphere conditions; rather, the high nasal cavity in humans was formed simply by evolutionary facial reorganization in the divergence of Homo from the other hominin lineages, impairing the air-conditioning performance. Even though the inhaled air is not adjusted well within the nasal cavity in humans, it can be fully conditioned subsequently in the pharyngeal cavity, which is lengthened in the flat-faced Homo. Thus, the air-conditioning faculty in the nasal passages was probably impaired in early Homo members, although they have survived successfully under the fluctuating climate of the Plio-Pleistocene, and then they moved "Out of Africa" to explore the more severe climates of Eurasia.

The higher the farther: distance-specific referential gestures in chimpanzees (Pan troglodytes).

Referential signals, such as manual pointing or deictic words, allow individuals to efficiently locate a specific entity in the environment, using distance-specific linguistic and/or gestural units. To explore the evolutionary prerequisites of such deictic ability, the present study investigates the ability of chimpanzees to adjust their communicative signals to the distance of a referent. A food-request paradigm in which the chimpanzees had to request a close or distant piece of food on a table in the presence/absence of an experimenter was employed. Our main finding concerns the chimpanzees adjusting their requesting behaviours to the distance of the food such that higher manual gestures and larger mouth openings were used to request the distant piece of food. To the best of our knowledge, this is the first study to demonstrate that chimpanzees are able to use distance-specific gestures.

Complete genome sequence of Streptococcus troglodytae TKU31 isolated from the oral cavity of a chimpanzee (Pan troglodytes).

Streptococcus troglodytae TKU31 was isolated from the oral cavity of a chimpanzee (Pan troglodytes) and was found to be the most closely related species of the mutans group streptococci to Streptococcus mutans. The complete sequence of TKU31 genome consists of a single circular chromosome that is 2,097,874 base pairs long and has a G + C content of 37.18%. It possesses 2082 coding sequences (CDSs), 65 tRNAs and five rRNA operons (15 rRNAs). Two clustered regularly interspaced short palindromic repeats, six insertion sequences and two predicted prophage elements were identified. The genome of TKU31 harbors some putative virulence associated genes, including gtfB, gtfC and gtfD genes encoding glucosyltransferase and gbpA, gbpB, gbpC and gbpD genes encoding glucan-binding cell wall-anchored protein. The deduced amino acid identity of the rhamnose-glucose polysaccharide F gene (rgpF), which is one of the serotype determinants, is 91% identical with that of S. mutans LJ23 (serotype k) strain. However, two other virulence-associated genes cnm and cbm, which encode the collagen-binding proteins, were not found in the TKU31 genome. The complete genome sequence of S. troglodytae TKU31 has been deposited at DDBJ/European Nucleotide Archive/GenBank under the accession no. AP014612.