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Individual differences in sulcal variation within the anterior and mid-cingulate cortex of the human brain, particularly the presence or absence of a paracingulate sulcus (PCGS), are associated with various motor and cognitive processes. Recently, it has been reported that chimpanzees possess a PCGS, previously thought to be a unique feature of the human brain. Here, we examined whether individual variation in the presence or absence of a PCGS as well as the variability in the intralimbic sulcus (ILS) are associated with oro-facial motor control, handedness for manual gestures, and sex in a sample of MRI scans obtained in 225 chimpanzees. Additionally, we quantified the depth of the cingulate sulcus (CGS) along the anterior-posterior axis and tested for association with oro-facial motor control, handedness, and sex. Chimpanzees with better oro-facial motor control were more likely to have a PCGS, particularly in the left hemisphere compared to those with poorer control. Male chimpanzees with better oro-facial motor control showed increased leftward asymmetries in the depth of the anterior CGS, whereas female chimpanzees showed the opposite pattern. Significantly, more chimpanzees had an ILS in the left compared to the right hemisphere, but variability in this fold was not associated with sex, handedness, or oro-facial motor control. Finally, significant population-level leftward asymmetries were found in the anterior portion of the CGS, whereas significant rightward biases were evident in the posterior regions. The collective results suggest that the emergence of a PCGS and enhanced gyrification within the anterior and mid-cingulate gyrus may have directly or indirectly evolved in response to selection for increasing oro-facial motor control in primates., (© The Author(s) 2021. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

It has been hypothesized that the evolution of tool use may have served as a preadaptation for the emergence of left hemispheric specialization in motor skill in humans. Here, we tested for intermanual differences in performance on a tool use task in a sample of 206 captive chimpanzees in relation to their sex, age, and hand preference. In addition, we examined heritability in tool use skill for the entire sample, as well as within 2 genetically isolated populations of captive chimpanzees. This was done to determine the degree of reproducibility in heritability on motor performance. The results revealed a significant effect of hand preference on intermanual differences in performance. Right-handed chimpanzees performed the task more quickly with their right compared with left hand. In contrast, no significant intermanual differences in performance were found in left- and ambiguous-handed apes. Tool use performance was found to be significantly heritable for overall performance, as well as separately for the left and right hands. Further, significant heritability in tool use performance was found in both populations of apes, suggesting these results were reproducible. The results are discussed in the context of evolutionary theories of handedness and hemispheric specialization and the genetic mechanisms that underlie their expression in primates, including humans. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

Investigations of behavioral lateralization in nonhuman primates yield important insights into brain-behavior relationships. In turn, they provide clues about both proximal and distal factors that shape the development and expression of association between motor asymmetries and underlying neural substrates. Nonhuman primates afford unique comparative opportunities to evaluate potential routes for the evolution of handedness, as well as to uncover relationships between behavioral lateralization and underlying neural, genetic, and physiological correlates. We examined hand preference in 22 rhesus monkeys and 79 chimpanzees using unimanual reaching tasks varying in postural stability and in a coordinated bimanual task. The majority of rhesus monkeys and chimpanzees showed significant lateral biases when reaching from a freestanding posture and when engaged in a coordinated bimanual task. Population-level directional bias was not evident for any task for rhesus monkeys and was observed only in the bimanual task for chimpanzees. We did not find consistent relationships between an individual's hand preference for different types of tasks. Both freestanding bipedal posture and coordinated bimanual hand use elicited significantly stronger lateral biases in reaching when compared with quadrupedal reaching. These data support the hypothesis that both degrees of postural instability and complex manipulation, such as bimanual coordination, may influence the expression of behavioral asymmetries in primates. These results demonstrate robust lateralization occurs at the individual level. Our results also highlight the need for greater consideration of task type and descriptive data in studies aimed at evaluating brain-behavior relationships and individual differences associated with hand preference. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

Increases brain size has been hypothesized to be inversely associated with the expression of behavioral and brain asymmetries within and between species. We tested this hypothesis by analyzing the relation between asymmetries in the planum temporale (PT) and different measures of the corpus callosum (CC) including surface area, streamline count as measured from diffusion tensor imaging, fractional anisotropy values and the ratio in the number of fibers to surface area in a sample of chimpanzees. We found that chimpanzees with larger PT asymmetries in absolute terms had smaller CC surface areas, fewer streamlines and a smaller ratio of fibers to surface area. These results were largely specific to male but not female chimpanzees. Our results partially support the hypothesis that brain asymmetries are linked to variation in corpus callosum morphology, although these associations may be sex-dependent., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

Chimpanzees are well known for their tool using abilities. Numerous studies have documented variability in tool use among chimpanzees and the role that social learning and other factors play in their development. There are also findings on hand use in both captive and wild chimpanzees; however, less understood are the potential roles of genetic and non-genetic mechanisms in determining individual differences in tool use skill and laterality. Here, we examined heritability in tool use skill and handedness for a probing task in a sample of 243 captive chimpanzees. Quantitative genetic analysis, based on the extant pedigrees, showed that overall both tool use skill and handedness were significantly heritable. Significant heritability in motor skill was evident in two genetically distinct populations of apes, and between two cohorts that received different early social rearing experiences. We further found that motor skill decreased with age and that males were more commonly left-handed than females. Collectively, these data suggest that though non-genetic factors do influence tool use performance and handedness in chimpanzees, genetic factors also play a significant role, as has been reported in humans., (© 2014 The Author(s) Published by the Royal Society. All rights reserved.)

There remain considerable questions regarding the evidence for population-level handedness in nonhuman primates when compared with humans. One challenge in comparing human and nonhuman primate handedness involves the procedures used to characterize individual handedness. Studies of human handedness use consistency in hand use within and between tasks as a basis for hand preference classification. In contrast, studies of handedness in nonhuman primates use statistical criteria for classifying handedness. In this study, we examined within- and between-task consistency in hand use as a means of characterizing individual handedness in a sample of 300 captive chimpanzees (Pan troglodytes). Chimpanzees showed population-level right-handedness for both within- and between-tasks consistency, though the proportion of right-handed chimpanzees was lower than what has typically been reported for humans. We further found that there were small, but significant, associations in hand use between measures. There were no significant sex or colony effects on the distribution of handedness. The results are discussed in the context of theories on the evolution of handedness in nonhuman primates.

Over the last century, the issue of brain lateralization in primates has been extensively investigated and debated, yet no previous study has reported eye preference in great apes. This study examined eye preference in 45 captive chimpanzees (Pan troglodytes) in response to various stimuli. Eye preference was assessed when animals looked through a hole that only accommodated one eye at an empty box, a mirror, a picture of a dog, a rubber snake, food biscuits, bananas, a rubber duck, and a video camera. Main effects of stimulus type were found for direction of eye preference, number of looks, and looking duration, but not for strength of eye preference. A left-eye bias was found for viewing the rubber snake and a right-eye bias was found for viewing the bananas, supporting theories that emotional valence may affect lateralized behaviors. In addition, a significant shift in eye preference took place from the initial look to subsequent looks when viewing the snake. These results are not consistent with previous reports of human eye preference and may reflect lateralization differences for emotional processing. No relationship between eye preference and previously recorded hand preference was found.

Functional and neuroanatomical asymmetries are an important characteristic of the human brain. The evolution of such specializations in the human cortex has provoked great interest in primate brain evolution. Most research on cortical sulci has revolved around linear measurements, which represent only one dimension of sulci organization. Here, we used a software program (BrainVISA) to quantify asymmetries in cortical depth and surface area from magnetic resonance images in a sample of 127 chimpanzees and 49 macaques. Population brain asymmetries were determined from 11 sulci in chimpanzees and seven sulci in macaques. Sulci were taken from the frontal, temporal, parietal, and occipital lobes. Population-level asymmetries were evident in chimpanzees for several sulci, including the fronto-orbital, superior precentral, and sylvian fissure sulci. The macaque population did not reveal significant population-level asymmetries, except for surface area of the superior temporal sulcus. The overall results are discussed within the context of the evolution of higher order cognition and motor functions., (Copyright © 2012 Elsevier Inc. All rights reserved.)

Whether lateralization of communicative signalling in non-human primates might constitute prerequisites of hemispheric specialization for language is unclear. In the present study, we examined (i) hand preference for a communicative gesture (clapping in 94 captive chimpanzees from two research facilities) and (ii) the in vivo magnetic resonance imaging brain scans of 40 of these individuals. The preferred hand for clapping was defined as the one in the upper position when the two hands came together. Using computer manual tracing of regions of interest, we measured the neuroanatomical asymmetries for the homologues of key language areas, including the inferior frontal gyrus (IFG) and planum temporale (PT). When considering the entire sample, there was a predominance of right-handedness for clapping and the distribution of right- and left-handed individuals did not differ between the two facilities. The direction of hand preference (right- versus left-handed subjects) for clapping explained a significant portion of variability in asymmetries of the PT and IFG. The results are consistent with the view that gestural communication in the common ancestor may have been a precursor of language and its cerebral substrates in modern humans.

Whether or not nonhuman primates exhibit population-level handedness remains a topic of considerable scientific debate. Here, we examined handedness for coordinated bimanual actions in a sample of 777 great apes including chimpanzees, bonobos, gorillas, and orangutans. We found population-level right-handedness in chimpanzees, bonobos and gorillas, but left-handedness in orangutans. Directional biases in handedness were consistent across independent samples of apes within each genus. We suggest that, contrary to previous claims, population-level handedness is evident in great apes but differs among species as a result of ecological adaptations associated with posture and locomotion. We further suggest that historical views of nonhuman primate handedness have been too anthropocentric, and we advocate for a larger evolutionary framework for the consideration of handedness and other aspects of hemispheric specialization among primates., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

Although behavioral lateralization is known to correlate with certain aspects of brain asymmetry in primates, there are limited data concerning hemispheric biases in the microstructure of the neocortex. In the present study, we investigated whether there is asymmetry in synaptophysin-immunoreactive puncta density and protein expression levels in the region of hand representation of the primary motor cortex in chimpanzees (Pan troglodytes). Synaptophysin is a presynaptic vesicle-associated protein found in nearly all synapses of the central nervous system. We also tested whether there is a relationship between hand preference on a coordinated bimanual task and the interhemispheric distribution of synaptophysin as measured by both stereologic counts of immunoreactive puncta and by Western blotting. Our results demonstrated that synaptophysin-immunoreactive puncta density is not asymmetric at the population level, whereas synaptophysin protein expression levels are significantly higher in the right hemisphere. Handedness was correlated with interindividual variation in synaptophysin-immunoreactive puncta density. As a group, left-handed and ambidextrous chimpanzees showed a rightward bias in puncta density. In contrast, puncta densities were symmetrical in right-handed chimpanzees. These findings support the conclusion that synapse asymmetry is modulated by lateralization of skilled motor behavior in chimpanzees.

It has been proposed that human right handedness is determined by genetic factors associated with the emergence of language, whereas non-human primate handedness is determined by random, non-genetic factors. These different mechanisms account for differences in the distribution of handedness between human and non-human primates. Here we report evidence that genetic factors play a role in the determination of handedness in chimpanzees. We further report that differential rearing has no influence on the expression of handedness in related individuals. Contrary to many theories of the origin of handedness, these results indicate that genetic factors have a significant influence on handedness in chimpanzees.

Humans throw right-handed, and it has been suggested that the neurophysiological demands of aimed throwing may have served as a precursor to the evolution of left hemisphere specialization for linguistic functions. Although there are descriptions of throwing by wild and captive chimpanzees (Pan troglodytes), systematic observations of aimed throwing and handedness have not been reported. In this article, evidence of population-level right-handedness for throwing is reported in 2 samples of captive chimpanzees. It is further reported that right-handed throwing is more pronounced than other measures of handedness in captive chimpanzees. The implications of these findings are discussed in the context of theories relating throwing to the evolution of lateralization for language functions., (((c) 2005 APA, all rights reserved).)

This article describes the distribution and development of handedness for manual gestures in captive chimpanzees. Data on handedness for unimanual gestures were collected in a sample of 227 captive chimpanzees. Handedness for these gestures was compared with handedness for three other measures of hand use: tool use, reaching, and coordinated bimanual actions. Chimpanzees were significantly more right-handed for gestures than for all other measures of hand use. Hand use for simple reaching at 3 to 4 years of age predicted hand use for gestures 10 years later. Use of the right hand for gestures was significantly higher when gestures were accompanied by a vocalization than when they were not. The collective results suggest that left-hemisphere specialization for language may have evolved initially from asymmetries in manual gestures in the common ancestor of chimpanzees and humans, rather than from hand use associated with other, non-communicative motor actions, including tool use and coordinated bimanual actions, as has been previously suggested in the literature.

Population-level right-handedness has historically been considered a hallmark of human evolution. Even though recent studies in chimpanzees (Pan troglodytes) have demonstrated population-level right-handedness for certain behaviors, some have questioned the validity and consistency of these findings by arguing that reported laterality effects are specific to certain colonies of apes and to those chimpanzees reared by humans. The authors report evidence of population-level right-handedness in 3 separate colonies of chimpanzees. Moreover, handedness in the 3 colonies was unrelated to the proportion of subjects that were raised by humans. This is the strongest evidence to date that population-level handedness is evident in chimpanzees and is not an artifact of human rearing., (((c) 2004 APA, all rights reserved))