Multiple measures of hand-use lateralization in the ring-tailed lemur (Lemur catta)

Evaluated a free-ranging matriline of 13 ring-tailed lemurs (Lemur catta) from videotaped records for lateralized hand use with 2 tasks and 4 measures: food reaching, feeding posture, duration of food holding, and manipulation of food between mouth and hand while eating. Binomial z scores determined 7 lemurs to be left preferent in reaching, 3 right, and 3 ambipreferent. Ideographic analyses suggested possible sex-linked and early experience twin effects. When compared to right and ambipreferent lemurs, left reach preferent lemurs used the left hand more but bimanuals grasped less in food holding and also engaged in less hand-mouth food manipulation. The tendency to manipulate food was not correlated with bimanual holding but was inversely related to left hand holding and directly related to right hand holding. These patterns are discussed as possible precursors of human bimanual manipulation.     

Implicit sequence learning in ring‐tailed lemurs (Lemur catta)

Implicit learning involves picking up information from the environment without explicit instruction or conscious awareness of the learning process. In nonhuman animals, conscious awareness is impossible to assess, so we define implicit learning as occurring when animals acquire information beyond what is required for successful task performance. While implicit learning has been documented in some nonhuman species, it has not been explored in prosimian primates. Here we ask whether ring‐tailed lemurs (Lemur catta) learn sequential information implicitly. We tested lemurs in a modified version of the serial reaction time task on a touch screen computer. Lemurs were required to respond to any picture within a 2 × 2 grid of pictures immediately after its surrounding border flickered. Over 20 training sessions, both the locations and the identities of the images remained constant and response times gradually decreased. Subsequently, the locations and/or the identities of the images were disrupted. Response times indicated that the lemurs had learned the physical location sequence required in original training but did not learn the identity of the images. Our results reveal that ring‐tailed lemurs can implicitly learn spatial sequences, and raise questions about which scenarios and evolutionary pressures give rise to perceptual versus motor‐implicit sequence learning.     

Analyses of feeding lateralization in the small-eared bushbaby (Otolemur garnettii): a comparison with the ring-tailed lemur (Lemur catta)

Feeding related lateralization was examined in a population of 23 small-eared bushbabies (Otolemur garnettii). The three measures used to determine lateralization were food reaching, holding, and manipulation. Sex and age differences were found, with adult females showing a strong right bias and adult males a left bias. Juvenile males were weakly lateralized and less consistent across measures than adult animals. The use of standard scores to assess lateralization allowed species comparisons to be made. The results of this study were compared with results from a previous study on lateralization in the ring-tailed lemur (Lemur catta). Species comparisons found sex differences to be a stronger factor in lateralization than species differences.     

Ring-tailed lemurs (Lemur catta) exploit information about what others can see but not what they can hear

Studies suggest that haplorhine primates are sensitive to what others can see and hear. Using two experimental designs, we tested the hypothesis that ring-tailed lemurs (N = 16) are also sensitive to the visual and auditory perception of others. In the first task, we used a go/no–go design that required lemurs to exploit only auditory information. In the second task, we used a forced-choice design where lemurs competed against a human who would prevent them from obtaining food if their approaches were detected. Subjects were given the choice of obtaining food silently or noisily when the competitor’s back was turned. They were also given the choice to obtain food when the competitor could either see them or not. Here, we replicate the findings of previous studies indicating that ring-tailed lemurs are sensitive to whether they can be seen; however, we found no evidence that subjects are sensitive to whether others can hear them. Our findings suggest that ring-tailed lemurs converge with haplorhine primates only in their sensitivity to the visual information of others. The results emphasize the importance of investigating social cognition across sensory domains in order to elucidate the cognitive mechanisms that underlie apparently complex social behavior. These findings also suggest that the social dynamics of haplorhine groups impose greater cognitive demands than lemur groups, despite similarities in total group size.     

A comparative analysis of serial and free recall

Multitrial free and serial recall tasks differ both in recall instruction and in presentation order across trials. Waugh (1961) compared these paradigms with an intermediate condition: free recall with constant presentation order. She concluded that differences between free and serial recall were due only to recall instructions, and not to presentation order. The present study reevaluated the relation between free and serial recall, using Waugh's three conditions. By examining recall transitions and the organization of information retained across trials, we conclude that presentation order is an important factor, causing participants to exhibit the same temporal associations in serial recall and in free recall with constant presentation order.     

How prosimian primates represent tools: experiments with two lemur species (Eulemur fulvus and Lemur catta)

The authors examined how 2 lemur species (Eulemur fulvus and Lemur catta) reason about tools. Experiment 1 allowed subjects to use 1 of 2 canes to retrieve an inaccessible food reward. Lemurs learned to solve this problem as quickly as other primates. Experiment 2 then presented subjects with novel tools differing from the originals along one featural dimension. Subjects attended more to tools' sizes than to their colors and made no distinction between tools' shapes and textures. Experiments 3 and 4 presented problems in which some of the tools' orientations had to be modified relative to the food. Subjects performed well on these problems, sometimes modifying the position of the tool. These results are discussed in light of the performance of other primates on this task.     

Expectations about numerical events in four lemur species (Eulemur fulvus, Eulemur mongoz, Lemur catta and Varecia rubra)

Although much is known about how some primates—in particular, monkeys and apes—represent and enumerate different numbers of objects, very little is known about the numerical abilities of prosimian primates. Here, we explore how four lemur species (Eulemur fulvus, E. mongoz, Lemur catta, and Varecia rubra) represent small numbers of objects. Specifically, we presented lemurs with three expectancy violation looking time experiments aimed at exploring their expectations about a simple 1+1 addition event. In these experiments, we presented subjects with displays in which two lemons were sequentially added behind an occluder and then measured subjects duration of looking to expected and unexpected outcomes. In experiment 1, subjects looked reliably longer at an unexpected outcome of only one object than at an expected outcome of two objects. Similarly, subjects in experiment 2 looked reliably longer at an unexpected outcome of three objects than at an expected outcome of two objects. In experiment 3, subjects looked reliably longer at an unexpected outcome of one object twice the size of the original than at an expected outcome of two objects of the original size. These results suggest that some prosimian primates understand the outcome of simple arithmetic operations. These results are discussed in light of similar findings in human infants and other adult primates.This revised version was published online in March 2005 with corrections to Fig. 6.     

An endogenous distributed model of ordering in serial recall

We introduce a distributed model of memory for serial order, called SOB, that produces ordered serial recall by relying on encoding and retrieval processes that are endogenous to the model. SOB explains the basic shape of the serial position curve, the pattern of errors during recall (including the balance between transpositions, omissions, intrusions, and erroneous repetitions), the effects of list length on the distribution of errors, the overall level of recall and response latency, and the effects of natural language frequency on recall performance. In addition, contrary to several recent suggestions, SOB demonstrates that distributed representations can support unambiguous recall, selective response suppression, and novelty-sensitive encoding.     

Spontaneous social orienting and gaze following in ringtailed lemurs (<Emphasis Type="Italic">Lemur catta</Emphasis>)

Both human and nonhuman primates preferentially orient toward other individuals and follow gaze in controlled environments. Precisely where any animal looks during natural behavior, however, remains unknown. We used a novel telemetric gaze-tracking system to record orienting behavior of ringtailed lemurs (Lemur catta) interacting with a naturalistic environment. We here provide the first evidence that ringtailed lemurs, group-living prosimian primates, preferentially gaze towards other individuals and, moreover, follow other lemurs’ gaze while freely moving and interacting in naturalistic social and ecological environments. Our results support the hypothesis that stem primates were capable of orienting toward and following the attention of other individuals. Such abilities may have enabled the evolution of more complex social behavior and cognition, including theory of mind and language, which require spontaneous attention sharing. This is the first study to use telemetric eye-tracking to quantitatively monitor gaze in any nonhuman animal during locomotion, feeding, and social interaction. Moreover, this is the first demonstration of gaze following by a prosimian primate and the first to report gaze following during spontaneous interaction in naturalistic social environments. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

A computer analysis of serial interactions in spaced responding

Serial dependencies in interresponse times were studied by means of a digital computer. In monkeys exposed to a DRL 20-sec schedule of reinforcement, serial interactions appeared at all stages of training. Early in training the serial effects consisted of trains of relatively long interresponse times interspersed among trains of relatively short ones. Later on, the serial effects appeared to be characterized by a tendency to drift up and down in long wavelength periods around the minimum interval required for reinforcement. After training to a point at which most interresponse times produced reinforcement, serial effects of a still more subtle nature appeared. These effects were made apparent by autocorrelation and power spectrum methods and consisted of both long-term and extremely short-term fluctuations in interresponse times.     

Representation of serial order in pigeons (Columba livia)

In Experiment 1, 2 groups of pigeons were trained to respond to either a 4-item (A→B→C→D) or 5-item (A→B→C→D→E) list. After learning their respective list, half of the subjects were trained on a positive pair with reinforcement provided when pairs were responded to in the order true to that of the original sequence (4-item: B→C; 5-item: B→D). The remaining subjects were trained on a negative pair with reinforcement provided for responding to the pairs in the order opposite to that learned in the original sequence (4-item: C→B; 5-item: D→B). Subjects in the positive pair condition learned their respective pair faster than did subjects in the negative pair condition. In Experiment 2, after reaching criterion on a 4-item list, subjects received 16 BC probe trials spread across 4 sessions of training. Subjects performed significantly above chance on the probe trials. The performance of our subjects in Experiments 1 and 2 demonstrates that, similar to monkeys, pigeons form a representation of the lists that they learn.     

Representation of serial order in monkeys (Cebus apella)

Cebus monkeys were trained on a five-item serial learning task, symbolized as ABCDE; the initial stages of training were on the shorter subseries AB, ABC, and ABCD. To assess the monkeys' knowledge of the sequential position of each item, pair-wise tests were given to 2 subjects after acquisition of the ABCD series and to 4 subjects after reaching criterion on the ABCDE series. In both tests, the monkeys performed at high levels on the interior pairs, which were BC for the ABCD series, and BC, BD, and CD for the ABCDE series. These results, as well as the orderly relations observed in the pair-wise tests between first-item response latency and first-item position and between second-item response latency and number of missing items, indicated that the monkeys had developed a well-organized internal representation of the four- and five-item series. Although pigeons are also capable of learning four-item and five-item series, they apparently do not develop a comparable representational structure. The disparity between the monkeys' and pigeons' representational competence for serial order is predictable from the difference in their capacities for associative transitivity.     

A comparative analysis of global and local processing of hierarchical visual stimuli in young children (Homo sapiens) and monkeys (Cebus apella)

Results obtained with preschool children (Homo sapiens) were compared with results previously obtained from capuchin monkeys (Cebus apella) in matching-to-sample tasks featuring hierarchical visual stimuli. In Experiment 1, monkeys, in contrast with children, showed an advantage in matching the stimuli on the basis of their local features. These results were replicated in a 2nd experiment in which control trials enabled the authors to rule out that children used spurious cues to solve the matching task. In a 3rd experiment featuring conditions in which the density of the stimuli was manipulated, monkeys' accuracy in the processing of the global shape of the stimuli was negatively affected by the separation of the local elements, whereas children's performance was robust across testing conditions. Children's response latencies revealed a global precedence in the 2nd and 3rd experiments. These results show differences in the processing of hierarchical stimuli by humans and monkeys that emerge early during childhood.     

A qualitative and serial analysis of social problem solving in aggressive boys

Social problem solving skills in 84 elementary school aged boys were assessed to identify those responses most salient in the prediction of ratings of aggressive and socially withdrawn behavior. Aggressive and socially withdrawn boys did not differ from the control group in the number of effective first solutions generated to the hypothetical stories. The control group generated significantly more effective solutions as second alternatives to the stories than did the aggressive or socially withdrawn boys. In addition, the number of effective second responses generated significantly predicted both aggression and social withdrawal after controlling for verbal problem solving. Results are discussed in terms of differences in cognitive processes as well as implications for treatment.This research was partially supported by a grant to the first author from the Charles Rieley Armington Foundation on Values in Children. The authors wish to thank Dennis Drotar, Douglas Detterman, and Met Evans for their assistance with this project.