Comparative assessment of handedness for a coordinated bimanual task in chimpanzees (Pan troglodytes), gorillas (Gorilla gorilla) and orangutans (Pongo pygmaeus)

Hand preferences for a coordinated bimanual task were assessed in a sample of 31 captive gorillas (Gorilla gorilla) and 19 captive orangutans (Pongo pygmaeus) and were compared with chimpanzee (Pan troglodytes) hand preferences in subjects that were matched on the basis of age, sex, and rearing history. The task required that the apes remove food from the inside edges of a symmetrical polyvinyl chloride pipe presented to them in their home cages. The results indicate significant species differences with chimpanzees showing population-level right-handedness and orangutans showing population-level left-handedness. The gorillas showed a nonsignificant trend toward right-handedness. The results are discussed in terms of possible ecological or biomechanical factors that may influence hand preferences in different ape species.     

Response of captive lowland gorillas (Gorilla gorilla gorilla) to different housing conditions: testing the aggression-density and coping models

Several studies have suggested that primates react differently to spatial reduction. In this article, the authors tested some general hypotheses on primate response to spatial reduction by studying the Apenheul lowland gorillas (Gorilla gorilla gorilla; Apeldoorn, the Netherlands). The frequency of conflicts did not greatly change between the 2 housing settings, thus not supporting the density- aggression model. Indoor, gorillas performed touching behavior more often and increased their level of reconciliation. These findings support the coping model. Indoor, the gorillas also maintained broader interindividual distances by increasing the levels of sitting alone, avoidance, and dismissing behaviors. In conclusion, the Apenheul gorillas modified selectively the distribution of some patterns typical of their behavioral repertoire in response to a high-density condition. Both avoidance and coping tactics were used, thus revealing high levels of behavioral flexibility in this species.     

Color preferences in gorillas (Gorilla gorilla gorilla) and chimpanzees (Pan troglodytes)

Color plays an important biological role in the lives of many animals, with some species exhibiting preferences for certain colors over others. This study explored the color preferences of two species of ape, which, like humans, possess trichromatic color vision. Six western lowland gorillas, and six chimpanzees, housed in Belfast Zoological Gardens, were exposed to three stimuli (cloths, boxes, sheets of acetate) in red, blue, and green. Six stimuli of the same nature, in each of the three colors, were provided to both species for 5 days per stimulus. The amount of interest that the animals showed toward each stimulus of each color was recorded for 1 hr. Results showed that the apes, both when analyzed as two separate groups, and when assessed collectively, showed significant color preferences, paying significantly less attention to the red-, than to the blue- or green-colored stimuli. The animals' interest in the blue- and green-colored stimuli did not differ significantly. Overall, the findings suggest that gorillas and chimpanzees, our closest living relatives, may harbor color preferences comparable to those of humans and other species.     

Quantity estimation and comparison in western lowland gorillas (Gorilla gorilla gorilla)

We investigated the quantity judgment abilities of two adult male western lowland gorillas (Gorilla gorilla gorilla) by presenting discrimination tasks on a touch-screen computer. Both gorillas chose the larger quantity of two arrays of dot stimuli. On some trials, the relative number of dots was congruent with the relative total area of the two arrays. On other trials, number of dots was incongruent with area. The gorillas were first tested with static dots, then with dots that moved within the arrays, and finally on a task where they were required to discriminate numerically larger subsets within arrays of moving dots. Both gorillas achieved above-chance performance on both congruent and incongruent trials with all tasks, indicating that they were able to use number as a cue even though ratio of number and area significantly controlled responding, suggesting that number was not the only relevant dimension that the gorillas used. The pattern of performance was similar to that found previously with monkeys and chimpanzees but had not previously been demonstrated in gorillas within a computerized test format, and with these kinds of visual stimuli.     

Imitative learning by captive western lowland gorillas (Gorilla gorilla gorilla) in a simulated food-processing task

Although field studies have suggested the existence of cultural transmission of foraging techniques in primates, identification of transmission mechanisms has remained elusive. To test experimentally for evidence of imitation in the current study, we exposed gorillas (Gorilla gorilla gorilla) to an artificial fruit foraging task designed by A. Whiten and D. M. Custance (1996). Gorillas (n=6) watched a human model remove a series of 3 defenses around a fruit. Each of the defenses was removed using 1 of 2 alternative techniques. Subsequent video analysis of gorillas' behavior showed a significant tendency to copy the observed technique on 1 of the individual defenses and the direction of removal on another defense. This is the first statistically reliable evidence of imitation in gorillas. Sequence of defense removal was not replicated. The gorillas' responses were most similar to those of chimpanzees.     

Target animacy influences gorilla handedness

We investigated the unimanual actions of a biological family group of twelve western lowland gorillas (Gorilla gorilla gorilla) using a methodological approach designed to assess behavior within social context from a bottom-up perspective. Measures of both the lateralization of unimanual actions (left, right) and the target of the action (animate, inanimate) were assessed during dual, synchronized video observations of naturalistic behavior. This paper demonstrates a corelationship between handedness and the animate quality of the target object. Analyses demonstrated a significant interaction between lateralized unimanual actions and target animacy and a right-hand bias for actions directed toward inanimate targets. We suggest that lateralized motor preference reflects the different processing capabilities of the left and right hemispheres, as influenced by the emotive (animate) and/or functional (inanimate) characteristics of the target, respectively.     

Spontaneously occurring mother-infant swapping and the relationships of infants with their biological and foster mothers in a captive group of lowland gorillas (Gorilla gorilla gorilla)

The present study describes spontaneously occurring infant-mother swapping and the relationships of infants with their biological and foster mothers after swapping in a captive social group of lowland gorillas (Gorilla gorilla gorilla). After infant-swapping took place between a primiparous mother with a neonate and a multiparous mother with a 10-month-old infant, the mothers provided appropriate maternal care, including nursing, to their adopted infants, but the older infant still sought refuge with its biological mother more frequently than with its foster mother. Almost 2 years later, the younger infant who was weaned by its foster mother began nursing from its biological mother; the latter subsequently weaned her adopted infant. Thereafter, the older infant who was weaned exhibited an abnormal hair-plucking behavior. These results indicate that it was psychologically stressful for the gorilla infant to quickly transfer its attachment figure from the mother to another adult female when the mother continued to remain in the group.     

Mother-infant interactions in western Lowland Gorillas (Gorilla gorilla gorilla): spatial relationships,communication, and opportunities for social learning

This study investigated mother-infant interactions in lowland gorillas (Gorilla gorilla gorilla) with particular focus on the relative role of mothers and infants in creating situations that are potentially conducive to infant social learning. Eleven gorilla mother-infant dyads were focally observed in weekly 1-hr sessions for 12 months. Spatial relationships were affected by age as well as by ambient temperature. Although the youngest infant was encouraged by its mother to walk and climb, mothers showed little or no encouragement in other contexts. In contrast, infants were quite interested in their mothers' activities, on some occasions repeated their mother's behavior, and actively encouraged their mothers to share food, play, or follow them. These findings suggest that gorilla infants are more active than their mothers in creating situations that are potentially conducive to the acquisition of knowledge or skills.     

Gestural communication of the gorilla (Gorilla gorilla): repertoire, intentionality and possible origins

Social groups of gorillas were observed in three captive facilities and one African field site. Cases of potential gesture use, totalling 9,540, were filtered by strict criteria for intentionality, giving a corpus of 5,250 instances of intentional gesture use. This indicated a repertoire of 102 gesture types. Most repertoire differences between individuals and sites were explicable as a consequence of environmental affordances and sampling effects: overall gesture frequency was a good predictor of universality of occurrence. Only one gesture was idiosyncratic to a single individual, and was given only to humans. Indications of cultural learning were few, though not absent. Six gestures appeared to be traditions within single social groups, but overall concordance in repertoires was almost as high between as within social groups. No support was found for the ontogenetic ritualization hypothesis as the chief means of acquisition of gestures. Many gestures whose form ruled out such an origin, i.e. gestures derived from species-typical displays, were used as intentionally and almost as flexibly as gestures whose form was consistent with learning by ritualization. When using both classes of gesture, gorillas paid specific attention to the attentional state of their audience. Thus, it would be unwarranted to divide ape gestural repertoires into ‘innate, species-typical, inflexible reactions’ and ‘individually learned, intentional, flexible communication’. We conclude that gorilla gestural communication is based on a species-typical repertoire, like those of most other mammalian species but very much larger. Gorilla gestures are not, however, inflexible signals but are employed for intentional communication to specific individuals.     

Development of hand preferences in marmosets (Callithrix jacchus) and effects of aging

The authors investigated the development of unimanual hand use and hand preferences during feeding in 15 marmosets (Callithrix jacchus), ages birth to 51-70 months. Bimanual hand use was common at 1-2 months, but by 5-8 months unimanual holding had developed and so had significant hand preferences. Half of the marmosets preferred to pick up and take food to the mouth with the left hand, and half preferred the right hand. Individuals maintained the same hand preference at all ages examined. Significant relationships were also found between the postures adopted during feeding and the direction of hand preferences displayed by juvenile marmosets. There was a positive correlation between increased suspension and increased left-hand preference, and a negative correlation between increased feeding in a tripedal posture and increased left-hand preference. These results are discussed in terms of motor development and hemispheric specialization.     

Relative numerousness judgment and summation in young and old Western lowland gorillas

The relationship between age, relative numerousness judgment, and summation was investigated in 11 Western lowland gorillas (Gorilla gorilla gorilla). Experiments 1 and 2 evaluated the gorillas' ability to select the larger of 2 food quantities before and with training. The majority of gorillas did not reliably select the larger quantity in Experiment 1 until receiving training to do so in Experiment 2. Experiment 3 evaluated their ability to select the larger of 2 pairs of quantities. All gorillas selected the larger pair more often than chance, and the old were less accurate and slower than were the young. For most gorillas, performance in relative numerousness judgment with training and summation was comparable with previous reports in chimpanzees and orangutans.     

Natural concepts in a juvenile gorilla (gorilla gorilla gorilla) at three levels of abstraction

The extent to which nonhumans are able to form conceptual versus perceptual discriminations remains a matter of debate. Among the great apes, only chimpanzees have been tested for conceptual understanding, defined as the ability to form discriminations not based solely on simple perceptual features of stimuli, and to transfer this learning to novel stimuli. In the present investigation, a young captive female gorilla was trained at three levels of abstraction (concrete, intermediate, and abstract) involving sets of photographs representing natural categories (e.g., orangutans vs. humans, primates vs. nonprimate animals, animals vs. foods). Within each level of abstraction, when the gorilla had learned to discriminate positive from negative exemplars in one set of photographs, a novel set was introduced. Transfer was defined in terms of high accuracy during the first two sessions with the new stimuli. The gorilla acquired discriminations at all three levels of abstraction but showed unambiguous transfer only with the concrete and abstract stimulus sets. Detailed analyses of response patterns revealed little evidence of control by simple stimulus features. Acquisition and transfer involving abstract stimulus sets suggest a conceptual basis for gorilla categorization. The gorilla's relatively poor performance with intermediate-level discriminations parallels findings with pigeons, and suggests a need to reconsider the role of perceptual information in discriminations thought to indicate conceptual behavior in nonhumans.     

Tracking the displacement of objects: a series of tasks with great apes (Pan troglodytes, Pan paniscus, Gorilla gorilla, and Pongo pygmaeus) and young children (Homo sapiens)

The authors administered a series of object displacement tasks to 24 great apes and 24 30-month-old children (Homo sapiens). Objects were placed under 1 or 2 of 3 cups by visible or invisible displacements. The series included 6 tasks: delayed response, inhibition test, A not B, rotations, transpositions, and object permanence. Apes and children solved most tasks performing at comparable levels except in the transposition task, in which apes performed better than children. Ape species performed at comparable levels in all tasks except in single transpositions, in which chimpanzees (Pan troglodytes) and bonobos (Pan paniscus) performed better than gorillas (Gorilla gorilla) and orangutans (Pongo pygmeaus). All species found nonadjacent trials and rotations especially difficult. The number of elements that changed locations, the type of displacement, and having to inhibit predominant reaching responses were factors that negatively affected the subjects' performance.     

Gorillas (Gorilla gorilla) and orangutans (Pongo pygmaeus) encode relevant problem features in a tool-using task

Two important elements in problem solving are the abilities to encode relevant task features and to combine multiple actions to achieve the goal. The authors investigated these 2 elements in a task in which gorillas (Gorilla gorilla) and orangutans (Pongo pygmaeus) had to use a tool to retrieve an out-of-reach reward. Subjects were able to select tools of an appropriate length to reach the reward even when the position of the reward and tools were not simultaneously visible. When presented with tools that were too short to retrieve the reward, subjects were more likely to refuse to use them than when tools were the appropriate length. Subjects were proficient at using tools in sequence to retrieve the reward.     

Differences in gorilla nettle-feeding between captivity and the wild: local traditions,species typical behaviors or merely the result of nutritional deficiencies?

Behavioral and cognitive studies on captive apes often pay little attention to the specific environmental conditions of their study subjects. A recent report form Byrne et al. (Anim Cogn doi:, 2011), comparing nettle-feeding techniques between captive and wild gorillas, claimed to document “the strongest evidence yet to come from any great ape that observational learning of a skilled conspecific” can allow social learning and culture in gorillas. An earlier study with similar findings placed emphasis instead on the many similarities and claims for species typical behavior, thus a genetic hypothesis instead of a cultural hypothesis. This commentary aims at formulating a third environmental hypothesis based on path-dependent behavioral differences owing to different diet and availability of nutritional resources of wild and captive gorillas. Captive diet provides gorillas with a much lower concentration of fibers. Gorillas are hindgut fermenters, and this deficit of natural fermentation of fibers may impact their health and their behavior in zoos. Results of Byrne et al.’s study will be discussed comparing feeding choice and availability of nutritional resources of wild and captive gorillas, showing that in captivity gorilla, motivation to consume certain food or certain plant parts may differ drastically from that of wild gorillas. This view does not intend to deny that social learning and culture may exist in gorillas, but to guide and encourage future works investigating social learning in great apes to take more accurately into account the living conditions and, when comparing populations, the possible environmental differences.     

Event memory and misinformation effects in a gorilla (<Emphasis Type="Italic">Gorilla gorilla gorilla</Emphasis>)

Event memory and misinformation effects were examined in an adult male gorilla ( Gorilla gorilla gorilla). The gorilla witnessed a series of unique events, involving a familiar person engaging in a novel behavior (experiment 1), a novel person engaging in a novel behavior (experiment 2), or the presentation of a novel object (experiment 3). Following a 5- to 10-min retention interval, a tester gave the gorilla three photographs mounted on wooden cards: a photograph depicting the correct person or object and two distractor photographs drawn from the same class. The gorilla responded by returning a photograph. If correct, he was reinforced with food. Across three experiments, the gorilla performed significantly above chance at recognizing the target photograph. In experiment 4, the gorilla showed at-chance performance when the event was followed by misinformation (a class-consistent, but incorrect photograph), but significantly above-chance performance when no misinformation occurred (either correct photograph or no photograph). Although the familiarity can account for these data, they are also consistent with an episodic-memory interpretation.     

Single-trial learning of "what" and "who" information in a gorilla (Gorilla gorilla gorilla): implications for episodic memory

Single-trial learning and long-term memory of "what" and "who" information were examined in an adult gorilla (Gorilla gorilla gorilla). We presented the gorilla with a to-be-remembered food item at the time of study. In Experiment 1, following a retention interval of either approximately 7 min or 24 h, the gorilla responded with one of five cards, each corresponding to a particular food. The gorilla was accurate on 70% of the short retention-interval trials and on 82% of the long retention-interval trials. In Experiment 2, the food stimulus was provided by one of two experimenters, each of whom was represented by a card. The gorilla identified the food (55% of the time) and the experimenter (82% of the time) on the short retention-interval trials. On the long retention-interval trials, the gorilla was accurate for the food (73%) and for the person (87%). The results are interpreted in light of theories of episodic memory. Electronic Publication     

The use of bouts and frequencies in the evaluation of hand preferences for a coordinated bimanual task in chimpanzees (Pan troglodytes): an empirical study comparing two different indices of laterality

Hand preferences for a coordinated bimanual task were assessed in 109 chimpanzees (Pan troglodytes). Hand preference was evaluated for 4 test sessions using bouts and frequencies of hand use to compare the sensitivity of each level of analysis in evaluating individual variation in handedness. Overall, significant population-level right-handedness was found using several different measures of hand use. Handedness indices based on bouts and frequencies were highly and significantly correlated. Moreover, hand preferences were consistent across tests despite efforts to situationally bias preference during each test. Taken together, these data do not support the view that bouts are a better level of analysis for evaluating hand preference. The results further suggest that hand preferences for coordinated bimanual actions are not influenced by situational factors and may reflect an inherent specialization of the left hemisphere for motor skill.     

Parasitoid wasps’ exposure to host-infested plant volatiles affects their olfactory cognition of host-infested plants

Using Cotesia vestalis, a parasitoid wasp of diamondback moth larvae and three crucifer plant species (cabbage, komatsuna, and Japanese radish), we examined the effects of exposure to host-infested plant volatiles from one plant species on a newly emerged wasp’s subsequent olfactory cognition of host-infested plant volatiles from the same or different plant species. The preference of C. vestalis between infested and uninfested plant volatiles was tested in a choice chamber. Volatile-inexperienced wasps significantly preferred infested cabbage and infested radish volatiles, but not infested komatsuna volatiles. After exposure to infested cabbage volatiles, wasps showed a significant preference for infested cabbage volatiles, while the significant preference for infested radish volatiles that had been observed in inexperienced wasps was no longer observed. After exposure to infested komatsuna volatiles, wasps significantly preferred infested komatsuna volatiles, and the pre-exposure significant preferences for infested cabbage volatiles and infested radish volatiles remained. After exposure to infested radish volatiles, the significant preferences for infested cabbage and infested radish volatiles did not change. Furthermore, wasps showed a significant preference for infested komatsuna volatiles. The compound compositions of the volatile blends from the three infested plant species were grouped separately in a principal coordinates analysis. The experience-based cognition of C. vestalis for host-infested plant volatiles of three plant species is discussed.     

Discrete quantity judgments in the great apes (Pan paniscus, Pan troglodytes, Gorilla gorilla, Pongo pygmaeus): the effect of presenting whole sets versus item-by-item

The authors examined quantity-based judgments for up to 10 items for simultaneous and sequential whole sets as well as for sequentially dropped items in chimpanzees (Pan troglodytes), gorillas (Gorilla gorilla), bonobos (Pan paniscus), and orangutans (Pongo pygmaeus). In Experiment 1, subjects had to choose the larger of 2 quantities presented in 2 separate dishes either simultaneously or 1 dish after the other. Representatives of all species were capable of selecting the larger of 2 quantities in both conditions, even when the quantities were large and the numerical distance between them was small. In Experiment 2, subjects had to select between the same food quantities sequentially dropped into 2 opaque cups so that none of the quantities were ever viewed as a whole. The authors found some evidence (albeit weaker) that subjects were able to select the larger quantity of items. Furthermore, the authors found no performance breakdown with the inclusion of certain quantities. Instead, the ratio between quantities was the best performance predictor. The authors conclude that quantity-based judgments rely on an analogical system, not a discrete object file model or perceptual estimation mechanism, such as subitizing.