Charles R. Menzel

Unprompted recall and reporting of hidden objects by a chimpanzee (Pan troglodytes) after extended delays.

The ability to recall features of environments not present to the senses is important in human thinking, planning, and communication, but to date there are almost no data on recall capabilities in nonverbal animals. In this study, the author used symbol knowledge as a tool to study chimpanzee memory. An 11-year-old female chimpanzee (Pan troglodytes) that had already learned a large number of arbitrarily designated geometric forms (lexigrams) watched as an experimenter hid an object in the woods outside her outdoor enclosure. The type and location of the object varied across trials. After an imposed delay of up to 16 h, the chimpanzee could interact indoors with a person who did not know that an object had been hidden, let alone the type or location of the object. A keyboard in the indoor cage displayed 256 lexigrams. From Trial 1, the chimpanzee seemed to do whatever it took to catch the persons attention and then touched the lexigram corresponding to the type of object hidden, pointed outdoors, went outdoors (if followed), and continued to point manually toward the object and vocalize until the person found the object. The subject indicated nonfood objects as well as more than 20 food types.

An efficient computerized testing method for the capuchin monkey (Cebus apella): adaptation of the LRC-CTS to a socially housed nonhuman primate species.

Even with advances in automated testing techniques, the capuchin monkey (Cebus apella) can be a difficult species to test in the laboratory, given its social/behavioral tendencies and typical activity pattern. Laboratories that maintain social colonies of capuchin monkeys are able to separate and test individuals, but the process can be very effortful and time consuming, and the resulting data can be modest in quantity. The present article describes procedures and apparatuses that were used to train a colony of computer-naive capuchin monkeys to quickly and reliably isolate themselves from group members and interact with a computerized test system in order to produce a large volume of data. Several elements that were important in motivating the monkeys to participate are discussed.

Bonobo (Pan paniscus) Spatial Memory and Communication in a 20-hectare Forest

We used an artificial language as a tool for the study of spatial memory organization in a young Pan paniscus. In the first experiment, we showed the bonobo a road sign just outside its indoor sleeping area. The sign indicated, by means of arbitrarily designated geometrical shapes (lexigrams), where food was hidden. Only 2 of the 15 locations were visible from the sign. Distances ranged up to 170 m from the sign. In 99 of 127 test trials the bonobo went directly to the designated location on its first move. In a second experiment, we presented the road sign at varied points in the woods rather than at the original fixed place. In these trials the goal was a preferred toy. The bonobos human companions were never told the location of the goal and distances were up to 650 m. In all 12 trials the bonobo led its companions to the designated place via an efficient path. The bonobo appeared to be able to move, based on the information provided by a lexigram, from almost any arbitrary starting location in its 20-ha environment to any one of the numerous goal locations.

Dissociation of cortisol and behavioral indicators of stress in an orangutan (Pongo pygmaeus) during a computerized task

Computerized testing can induce behavioral signs of frustration in apes. Three variations of a computer task were used to investigate the effects of inter-trial intervals and rate of cursor movement on frustrative behavior and cortisol in an orangutan. Behaviors were recorded during test sessions, and saliva was collected immediately after test sessions for cortisol assay. Behavioral results indicated that extended (20 sec) periods of delay between trials induced signs of frustration in the subject, including forceful manual manipulation of objects and self-scratching. However, cortisol results indicated that Hypothalamic-Pituitary-Adrenal (HPA) axis activity was not induced by task performance. Rather, cortisol levels were reduced during performance of computer tasks compared to baseline levels. Findings from this study suggest that behavioral and cortisol responses to stress induced by performance of computer testing can become dissociated. This study validates salivary cortisol as a measure of HPA activity in apes and demonstrates a normal circadian rhythm of cortisol release in an orangutan.

Memory and foraging theory: Chimpanzee utilization of optimality heuristics in the rank-order recovery of hidden foods

Many models from foraging theory and movement ecology assume that resources are encountered randomly. If food locations, types and values are retained in memory, however, search time could be significantly reduced, with concurrent effects on biological fitness. Despite this, little is known about what specific characteristics of foods, particularly those relevant to profitability, nonhuman animals can remember. Building upon previous observations, we hypothesized that chimpanzees (Pan troglodytes), after observing foods being hidden in a large wooded test area they could not enter, and after long delays, would direct (through gesture and vocalization) experimentally naïve humans to the reward locations in an order that could be predicted beforehand by the spatial and physical characteristics of those items. In the main experiment, various quantities of almonds, both in and out of shells and sealed in transparent bags, were hidden in the test area. The chimpanzees later directed searchers to those items in a nonrandom order related to quantity, shell presence/absence, and the distance they were hidden from the subject. The recovery sequences were closely related to the actual e/h profitability of the foods. Predicted recovery orders, based on the energetic value of almonds and independently-measured, individual-specific expected pursuit and processing times, were closely related to observed recovery orders. We argue that the information nonhuman animals possess regarding their environment can be extensive, and that further comparative study is vital for incorporating realistic cognitive variables into models of foraging and movement.

Spatial Memory and Monitoring of Hidden Items Through Spatial Displacements by Chimpanzees (Pan troglodytes).

This study examined chimpanzee (Pan troglodytes) short-term memory for food location in near space. In Experiments 1 and 2, either 1 or 2 items (chocolate pieces) were hidden in an array of 3 or 5 containers that either remained stationary or were rotated 180 degrees or 360 degrees. When the array remained stationary, the chimpanzees remembered both item locations. When arrays were rotated, however, chimpanzees found only 1 item. In Experiment 3, 2 items were hidden in an array of 7 cups. Both items were found at levels significantly better than chance. Ninety percent of errors were made after the 1st item was found, and errors reflected memory failure rather than a failure of inhibitory control.

Navigating two-dimensional mazes: Chimpanzees (Pan troglodytes) and capuchins (Cebus apella sp.) profit from experience differently

We examined whether navigation is impacted by experience in two species of nonhuman primates. Five chimpanzees (Pan troglodytes) and seven capuchin monkeys (Cebus apella) navigated a cursor, using a joystick, through two-dimensional mazes presented on a computer monitor. Subjects completed 192 mazes, each one time. Each maze contained one to five choices, and in up to three of these choices, the correct path required moving the cursor away from the Euclidean direction toward the goal. Some subjects completed these mazes in a random order (Random group); others in a fixed order by ascending number of choices and ascending number of turns away from goal (Ordered group). Chimpanzees in both groups performed equivalently, demonstrated fewer errors and a higher rate of self-correcting errors with increasing experience at solving the mazes, and made significantly fewer errors than capuchin monkeys. Capuchins were more sensitive to the mode of presentation than chimpanzees; monkeys in the Ordered group made fewer errors than monkeys in the Random group. However, capuchins’ performance across testing changed little, and they remained particularly susceptible to making errors when the correct path required moving away from the goal. Thus, these two species responded differently to the same spatial challenges and same learning contexts. The findings indicate that chimpanzees have a strong advantage in this task compared to capuchins, no matter how the task is presented. We suggest that differences between the species in the dynamic organization of attention and motor processes contribute to their differences in performance on this task, and predict similar differences in other tasks requiring, as this one does, sustained attention to a dynamic visual display and self-produced movements variably towards and away from a goal.

How young children and chimpanzees (Pan troglodytes) perceive objects in a 2D display: putting an assumption to the test

Object recognition research is typically conducted using 2D stimuli in lieu of 3D objects. This study investigated the amount and complexity of knowledge gained from 2D stimuli in adult chimpanzees (Pan troglodytes) and young children (aged 3 and 4 years) using a titrated series of cross-dimensional search tasks. Results indicate that 3-year-old children utilize a response rule guided by local features to solve cross-dimensional tasks. Four-year-old toddlers and adult chimpanzees use information about object form and compositional structure from a 2D image to guide their search in three dimensions. Findings have specific implications to research conducted in object recognition/perception and broad relevance to all areas of research and daily living that incorporate 2D displays.

How tufted capuchin monkeys (Cebus apella spp) and common chimpanzees (Pan troglodytes) align objects to surfaces: Insights into spatial reasoning and implications for tool use

This report addresses phylogenetic variation in a spatial skill that underlies tool use: aligning objects to a feature of a surface. Fragaszy and Cummins‐Sebrees [Behavioral and Cognitive Neuroscience Reviews 4:282–306, 2005] model of relational spatial reasoning and Skill Development and Perception–Action theories guided the design of the study. We examined how capuchins and chimpanzees place stick objects of varying shapes into matching grooves on a flat surface. Although most individuals aligned the long axis of the object with the matching groove more often than expected by chance, all typically did so with poor precision. Some individuals managed to align a second feature, and only one (a capuchin monkey) achieved above‐chance success at aligning three features with matching grooves. Our findings suggest that capuchins and chimpanzees do not reliably align objects along even one axis, and that neither species can reliably or easily master object placement tasks that require managing two or more spatial relations concurrently. Moreover, they did not systematically vary their behavior in a manner that would aid discovery of the affordances of the stick–surface combination beyond sliding the stick along the surface (which may have provided haptic information about the location of the groove). These limitations have profound consequences for the forms of tool use we can expect these individuals to master. Am. J. Primatol. 73:1012–1030, 2011.

Chimpanzees (Pan troglodytes) use markers to monitor the movement of a hidden item

Four chimpanzees (Pan troglodytes) monitored the movement of hidden items in arrays of opaque cups. A chocolate candy was hidden in an array of four cups and temporarily presented paper markers indicated the location of the candy (which otherwise was not visible). These markers were either non-symbolic or symbolic (lexigram) stimuli that in other contexts acted as a label for the hidden candy, and the array was either rotated 180° after the marker was removed or the array remained in the same location. For three of four chimpanzees, performance was better than chance in all conditions and there was no effect of the type of marker. These experiments indicate that chimpanzees can track the movement of a hidden item in an array of identical cups even when they never see the item itself, but only see a temporarily presented marker for the location of that item. However, there was no benefit to the use of symbolic as opposed to non-symbolic stimuli in this performance.