Analysis of lateralized components of feeding behavior in the ring-tailed lemur (Lemur catta).

Twenty-one ring-tailed lemurs (Lemur catta) were videotaped during feeding. They had previously been classified as left-, right- or ambipreferent on the basis of the hand used to reach for food. The feeding sequences provided duration-based measures of manipulation, hand and mouth lateralization, and posture during feeding sequences with 2 types of food. Hand preference for reaching and holding was stable over time and across measurement conditions. As a group, the lemurs grasped fruit bimanually more often than chow; however, right- and ambipreferent lemurs spent more time holding food bimanually than did left-preferent ones. Older animals consumed chow more quickly than did younger animals but did not differ in their rate of manipulation. Lemurs had a preference for using one side of the mouth in feeding, but this had no directional relation to preferred hand. Three postural feeding patterns were identified. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Hand preferences in unimanual and bimanual feeding by wild vervet monkeys (Cercopithecus aethiops).

Lateral preference was examined in spontaneous feeding actions in 2 troops of wild vervet monkeys (Cercopithecus aethiops). Processing of 4 foods (termites, leaf shoots, sugarcane, and fruit) was studied. Actions included unimanual reaching to moving objects, operating from an unstable posture, and coordinated bimanual processing. Between 19 and 31 subjects were available, according to the task. In 2 tasks, laterality of 2 independent stages was measured separately, giving 6 measures in all. On 4 of these measures, most monkeys were ambipreferent, and only a few showed significant hand preferences. Only for termite feeding and detaching material from fruits did the majority show significant lateralization; no tasks elicited exclusive use of 1 hand. Preference appeared labile, because in 2 tasks, population trends reversed with increasing age. No population trends to left or right were found; instead, these monkeys showed ambilaterality, with lateralization associated with task complexity. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Handedness as a function of sex and age in a large population of Lemur..

A population of 194 lemurs (Lemur spp.), 116 males and 78 females, from 1 to 30 years of age, was assessed for lateralized hand use in simple food reaching with a minimum of 100 reaches per animal. A hand preference was present in 80% of the population with a bias for use of the left hand that was most characteristic of male lemurs and young lemurs. The results confirm the presence of lateralization in prosimians, and we interpret the sex and age differences in relation to current theories of neural lateralization. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Visuospatial reaching preferences of common marmosets (Callithrix jacchus): An assessment of individual biases across a variety of tasks.

Using multiple measures of hand preference, the authors investigated lateralization at an individual level in 21 common marmosets. Despite showing group biases for sensory and communication functions, these same marmosets did not show a group bias in direction of lateralized hand use. Hand preferences were recorded on four novel reaching tasks requiring different levels of visual guidance and postural control. As found for simple food holding (with the same subjects), they displayed strong individual hand preferences but no group bias indicative of handedness. The strength of hand preference was influenced by task demands: stronger preferences were expressed when subjects adopted a suspended posture, and when "successful" versus "unsuccessful" foraging strategies were compared. Comparisons between visuospatial reaching and simple food holding preferences also revealed that half of the subjects displayed a division of function between the hands/hemispheres; subjects displayed opposing preferences in simple and visuospatial reaching, which would be beneficial for the performance of coordinated bimanual tasks. Given the apparent absence of a selective advantage for handedness, the authors suggest that hand preferences may reflect hemispheric dominance of other cognitive domains (i.e., temperament). (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Behavioral laterality in captive bonobos (Pan paniscus).

Behavioral laterality was studied in a captive group of 11 bonobos (Pan paniscus). In an observational method a significant left lateral bias in carrying and a right lateral bias in leading limb were found. Direction of lateral bias in carrying and leading limb was enhanced when Ss maintained a bipedal posture. Analysis of bimanual feeding behavior revealed a significant right-hand bias for eating when the Ss were holding food with their left hand. In an experimental method a significant shift toward greater right-hand use was found when Ss were required to maintain a bipedal, rather than a quadrupedal, posture. There was no evidence of lateral bias for measures of face touching, self-touching, or gestures. The results are discussed in the context of previous reports of primate laterality. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Bimanual interference associated with the selection of target locations.

Four experiments were conducted to identify the locus of interference observed during the preparation of bimanual reaching movements. Target locations were specified by color, and the right-hand and left-hand targets could be either the same or a different color. Movements of different amplitudes (Experiment 1) or different directions (Experiment 2) to targets of the same color were initiated more quickly than symmetric movements to targets of different colors. These results indicate that costs observed during bimanual movements arise during target selection rather than during motor programming. Experiments 3 and 4 further examined the interference associated with target selection. Reaction time costs were found with unimanual movements when the target was presented among distractors associated with responses for the other hand. Interference observed during bimanual reaching appears to reflect difficulty in segregating the response rules assigned to each hand. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Assessment of shoulder dimensions and angles of porcelain bonded to metal crown preparations

This research examined hand preference for a bimanual task in 45 tufted capuchin (Cebus apella) and 55 rhesus macaque (Macaca mulatta) monkeys. Investigators presented subjects with plastic tubes lined with food and noted which hand the animals used to hold the tubes and which hand the animals used to remove the food. Several significant findings emerged from this investigation. First, rhesus macaques, but not tufted capuchins, exhibited a population-level bias toward use of the right hand (although the difference in direction of hand preference between species was not significant). Second, capuchins exhibited greater hand preference strength than did macaques. Third, among capuchins, but not among macaques, hand preference strength was greater for adults than for immatures. Finally, both species used their index digit to remove food most frequently when compared with other digits. Findings of hand preference direction and strength in this study were compared with other findings noted for chimpanzees which performed a bimanual tube task in a previous study. The authors conclude that using the same procedure to compare hand preference across species represents a powerful research tool that can lead to a more complete understanding of the evolution and ontogenesis of primate handedness.     

Neuronal activity in the primate supplementary motor area and the primary motor cortex in relation to spatio-temporal bimanual coordination

Single neuronal activity was recorded from the supplementary motor area (SMA-proper and pre-SMA) and primary motor cortex (M1) in two Macaca fascicularis trained to perform a delayed conditional sequence of coordinated bimanual pull and grasp movements. The behavioural paradigm was designed to distinguish neuronal activity associated with bimanual coordination from that related to a comparable motor sequence but executed unimanually (left or right arm only). The bimanual and unimanual trials were instructed in a random order by a visual cue. Following the cue, there was a waiting period until presentation of a "go-signal", signalling the monkey to perform the instructed movement. A total of 143 task-related neurons were recorded from the SMA (SMA-proper, 62; pre-SMA, 81). Most SMA units (87%) were active in both unimanual contralateral and unimanual ipsilateral trials (bilateral neurons), whereas 9% of units were active only in unimanual contralateral trials and 3% were active only in unimanual ipsilateral trials. Forty-eight per cent of SMA task-related units were classified as bimanual, defined as neurons in which the activity observed in bimanual trials could not be predicted from that associated with unimanual trials when comparing the same events related to the same arm. For direct comparison, 527 neurons were recorded from M1 in the same monkeys performing the same tasks. The comparison showed that M1 contains significantly less bilateral neurons (75%) than the SMA, whereas the reverse was observed for contralateral neurons (22% in M1). The proportion of M1 bimanual cells (53%) was not statistically different from that observed in the SMA. The results suggest that both the SMA and M1 may contribute to the control of sequential bimanual coordinated movements. Interlimb coordination may then take place in a distributed network including at least the SMA and M1, but the contribution of other cortical and subcortical areas such as cingulate motor cortex and basal ganglia remains to be investigated.     

Alien hand syndrome: influence of neglect on the clinical presentation of frontal and callosal variants

Three patients with mesial frontal and extensive callosal lesions due to anterior cerebral artery infarction manifested an alien hand syndrome (AHS) with varied features. Patient 1 with left hemispheric lesion showed right hand's impulsive reaching and grasping and left hand's antagonistic movements to the right (intermanual conflict; IMC). Patients 2 and 3 with right hemispheric lesion manifested a left hemihypokinesia which was thought to have suppressed the frequency and amplitude or even the occurrence of left hand's reaching and grasping. IMC and other left hand's non-antagonistic, irrelevant movements to the right remained. Because the term "IMC" is often misused and not strictly defined, its association with right hand's reaching and grasping is quite uncommon, its significance as a sign of callosal disconnection is not well validated, and because left hand's reaching and grasping tend to be suppressed by motor neglect, a trend may then develop for the right hand to be the sole focus of pathological behaviour in patients with the so-called frontal AHS (Feinberg, Schindler, Flanagan et al., 1992).     

Posturally related variations in the hand preferences of the ruffed lemur (Varecia variegata variegata).

The hand preferences of 5 semi-free-ranging black-and-white ruffed lemurs were assessed by using three distinct testing procedures. Testing conditions varied in the extent to which they required animals to make a whole body postural adjustment prior to making a reach. Minimal bodily adjustment was necessary for free foraging, whereas discrete food presentations on land (DFP-land) and in a moat (DFP-moat) promoted a gross reorientation of the animal's entire body. In the DFP-moat condition 4 animals exhibited exclusive use of the left hand, and only 1 of 515 reaches was made with the right hand. Similarly, all 5 animals showed a pronounced left-hand preference in the DFP-land condition. The free-foraging condition revealed a hand preference for only 1 of the 5 subjects, and that preference was weak in comparison with those measured in the other two test conditions. These findings indicate that whole body postural adjustments critically influenced the expression of hand preference and should be taken into consideration in future studies of primate hand preferences. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Reach and posture hand preferences during arboreal feeding in sifakas (Propithecus sp.): A test of the postural origins theory of behavioral lateralization.

Sixteen sifakas (11 Propithecus verreauxi coquereli, 2 Propithecus verreauxi verreauxi form majori, and 3 Propithecus tattersalli) were videotaped as they fed on leaves in an arboreal context. The hand used to feed and the hand used to maintain postural stability was coded. For each subject, the lateral bias of the hand used to feed was opposite the hand used in postural support. Seven sifakas displayed no bias for feeding or posture-related hand use, 7 sifakas displayed significant feeding-related reach preferences for pulling branches to the mouth (5 left- and 2 right-hand preferences), and 9 sifakas exhibited significant hand preferences for postural support (2 left-, 7 right-hand preferent). Although these data do not strongly support the postural origins theory of behavioral lateralization, the modal preference pattern for sifakas that displayed significant hand preferences for posture and feeding involve a left bias for feeding and a right bias for postural support. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Are age trends in adult hand preference best explained by developmental shifts or generational differences?

Analyzed hand use in the performance of 6 everyday activities as a function of age in a sample of 654 adults (aged 18–85 yrs). An unskilled grasping task showed the largest shift toward consistent right-handedness as a function of age; skilled manipulation tasks, such as writing, exhibited weaker age-related trends. Analysis of individual preference patterns shows decreases in mixed right hand use and increases in consistent right hand use in older adults. There was also a weaker age-related reduction in mixed left hand use but the number of consistent left-handers remained a small, constant percentage across all age groups. Results are congruent with the predictions of 2 developmental hypotheses that stipulate a postulated age trend toward hand preference consistency, and a practice effect related to an environment arranged to favor right hand use. (French abstract) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

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. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Hand preference for a bimanual task in tufted capuchins (Cebus apella) and rhesus macaques (Macaca mulatta).

Examined hand preference for a bimanual task in 45 tufted capuchin (Cebus apella) and 55 rhesus macaque (Macaca mulatta) monkeys. Investigators presented subjects with plastic tubes lined with food and noted which hand the animals used to hold the tubes and which hand the animals used to remove the food. Several significant findings emerged from this investigation. First, rhesus macaques, but not tufted capuchins, exhibited a population-level bias toward use of the right hand (although the difference in direction of hand preference between species was not significant). Second, capuchins exhibited greater hand preference strength than did macaques. Third, among capuchins, but not among macaques, hand preference strength was greater for adults than for immatures. Finally, both species used their index digit to remove food most frequently when compared with other digits. Findings of hand preference direction and strength in this study were compared with other findings noted for chimpanzees which performed a bimanual tube task in a previous study. The authors conclude that using the same procedure to compare hand preference across species represents a powerful research tool that can lead to a more complete understanding of the evolution and ontogenesis of primate handedness. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Complexity in object manipulation by Japanese macaques (Macaca fuscata): A cross-sectional analysis of manual coordination in stone handling patterns.

Defined as a spontaneous stone-directed noninstrumental manipulative behavior, and comprised of multiple one-handed and (a)symmetrical/(un)coordinated two-handed patterns, stone handling (SH) is a good candidate for the study of complexity in object manipulation. We present a cross-sectional developmental analysis of SH complexity in Japanese macaques (Macaca fuscata), through the combined investigation of bimanuality, coordination, and symmetry in hand use. Bimanual SH patterns were more frequent than unimanual patterns. Among bimanual patterns, coordinated actions were more frequent than uncoordinated ones. We recorded five asymmetrical coordinated SH patterns with manual role differentiation, a form of hand use reminiscent of complex actions involving the use of tools in monkeys and apes. Bimanuality in SH was affected by body posture. Aging individuals performed less bimanual and less coordinated SH patterns than younger individuals. Our result on senescent males performing less bimanual patterns than senescent females was consistent with sex differences found in the late deterioration of complex manual movements in other species. Although some SH patterns represent a high degree of behavioral complexity, our results suggest that SH behavior is not as complex as tool-use or tool-manufacture in other nonhuman primates and hominids. (PsycINFO Database Record (c) 2011 APA, all rights reserved)     

Postural effects on manual reaching laterality in squirrel monkeys (Saimiri sciureus) and cotton-top tamarins (Saguinus oedipus).

Squirrel monkeys (Saimiri sciureus) and cotton-top tamarins (Saguinus oedipus) made unimanual food reaches from either a horizontal, quadrupedal posture or a vertical, clinging posture. No population-level handedness occurred in either species. However, in both species, directional lateral preferences weakly expressed for reaching from the stable quadrupedal posture were intensified in the vertical cling posture. This phenomenon, which is designated as soft handedness, may have been an evolutionary precursor to population-level handedness. Right or left turning by the squirrel monkeys before reaching closely predicted use of the right or left hand. However, the magnitude of the association decreased in the more highly lateralized vertical cling condition. This result suggests that as lateral hand preference increases, hand use may become increasingly independent of constraints from prior behavioral and environmental influences. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

A lateralized comparison of handedness and object proximity.

This study was motivated by the emerging hypothesis that right-handers are more strongly lateralized and perform better on various aspects of functional asymmetry than do left-handers. 84 Right- and 60 left-handers (undergraduates) were observed for hand selection responses to a unimanual task of reaching for a small cube in positions of right and left hemispace, prompting hemispheric decision-making related to hand dominance and attentional (visuospatial) stimuli. As predicted, left-handers did not respond with their preferred limb as consistently across positions as did right-handers. Additional inspection of the task suggests that being less lateralized may not be a disadvantage in this context, and that environmental influence may play a significant role in hand selection for a particular motor event. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Grip morphology and hand use in chimpanzees (Pan troglodytes): Evidence of a left hemisphere specialization in motor skill.

Three experiments on grip morphology and hand use were conducted in a sample of chimpanzees. In Experiment 1, grip morphology when grasping food items was recorded, and it was found that subjects who adopted a precision grip were more right-handed than chimpanzees using other grips. In Experiment 2, the effect of food type on grasping was assessed. Smaller food items elicited significantly more precision grips for the right hand. In Experiment 3, error rates in grasping foods were compared between the left and right hands. Significantly more errors were made for the left compared with the right hand. The cumulative results indicate that chimpanzees show a left-hemisphere asymmetry in motor skill that is associated with the use of precision grips. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Methodology of motor skill and laterality: new test of hand preference in Macaca nemestrina

Ten female pig-tailed macaques (Macaca nemestrina) were tested for hand preference and hand skill (i.e., speed of performance and error rate). The experimental task was naturalistic, calling for adaptive manual skills and fine manipulation: the monkeys had to remove small food rewards embedded in a vertical array, and precision opposition of thumb and forefinger was needed to extract each pellet. Each monkey was tested 10 times on 10 different days. The results indicated individual hand preference rather than population-level handedness; however, a tendency toward right predominance was found. The results on hand skill showed a relation between error rate and hand preference, as the preferred hand made fewer errors. A different and unexpected finding was obtained when skill was evaluated in terms of speed of performance: in adult subjects the left hand was quicker than the right. Therefore, different kinds of skill showed different patterns in relation to hand preference.     

Regional brain activity during tasks devoted to the central executive of working memory

Most previous PET studies investigating the central executive (CE) component of working memory found activation in the prefrontal cortex. However, the tasks used did not always permit to distinguish precisely the functions of the CE from the storage function of the slave systems. The aim of the present study was to isolate brain areas that subserve manipulation of information by the CE when the influence of storage function was removed. A PET activation study was performed with four cognitive tasks, crossing conditions of temporary storage and manipulation of information. The manipulation of information induced an activation in the right (BA 10/46) and left (BA 9/6) middle frontal gyrus and in the left parietal area (BA7). The interaction between the storage and manipulation conditions did not reveal any significant changes in activation. These results are in agreement with the hypothesis that CE functions are distributed between anterior and posterior brain areas, but could also reflect a simultaneous involvement of controlled (frontal) and automatic (parietal) attentional systems. In the other hand, the absence of interaction between the storage and manipulation conditions demonstrates that the CE is not necessarily related to the presence of a memory load.