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Research paper thumbnail of Hand Preferences for Bimanual Coordination in 77 Bonobos (Pan paniscus): Replication and Extension

International Journal of Primatology, Jan 11, 2011

The literature on manual laterality in nonhuman primates provides inconsistent and inconclusive f... more The literature on manual laterality in nonhuman primates provides inconsistent and inconclusive findings and is plagued by methodological issues (e.g., small samples, inconsistency in methods, inappropriate measures) and gaps. Few data are available on bonobos and these are only from small samples and for relatively simple tasks. We examined laterality in a large sample of bonobos for a complex task. We tested 48 bonobos from Lola Ya Bonobo sanctuary (DR Congo) in an extension of our previous study of 29 bonobos from 3 European zoos. We assessed hand preferences using the tube task, which involves bimanual coordination: one hand extracts food from a tube that is held by the other hand. This task is a good measure of laterality and it has been used in other studies. We recorded events (frequency) and independent bouts of food extraction. We found significant manual laterality, which was not influenced by the settings or rearing history. We observed little effect of sex and found an influence of age, with greater right hand use in adults. The laterality was marked, with strong preferences and most individuals being lateralized (when analyzing frequency). We found individual preferences, with no group-level bias, even when we combined the data from the sanctuary and the zoos to enlarge the sample to 77. These first data, for a complex task and based on a large sample, are consistent with previous findings in

Research paper thumbnail of Hand preference and its flexibility according to the position of the object: a study in cercopithecines examining spontaneous behaviour and an experimental task (the Bishop QHP task)

The extant literature on manual laterality in non-human primates is inconclusive, plagued by inco... more The extant literature on manual laterality in non-human primates is inconclusive, plagued by inconsistent or contradictory findings and by disturbing methodological issues (e.g. uncontrolled influential factors, comparability issues). The present study examined hand preference and its flexibility in 15 red-capped mangabeys (C. t. torquatus) and 13 Campbell's monkeys (C. c. campbelli), two species that differ in their degree of arboreality. We investigated the influence of the spatial position of the object on hand preference for reaching. We considered spontaneous behaviour (reaching for food during daily feeding) and an experimental task: the QHP task. The QHP is a task that is used in humans. This is a simple reaching task that involves high spatial constraints on hand use. In our study, the subject had to reach for items that were placed on a semi-circle in front of it on five positions, including in the centre position, in the ipsilateral space and in the contralateral space. We assessed hand preference for reaching in front (baseline condition), and we examined how this preference changed when reaching in lateral positions. For reaching in front, about half of the subjects were lateralized and no group-level bias occurred, for both spontaneous and experimental conditions. When considering reaching in the lateral positions, we observed that the position of the object influenced hand use: individuals used the hand that was closest to the object. The results are discussed in relation to previous findings in humans and in non-human primates and regarding theories on handedness and flexibility of hand preference.

Research paper thumbnail of The evolution of handedness in humans and great apes: a review and current issues

Population-level right-handedness is a defining characteristic of humans. Despite extensive resea... more Population-level right-handedness is a defining characteristic of humans. Despite extensive research, we still do not know the conditions or timing of its emergence in human evolution. We present a review of research into the origins of handedness, based on fossil and archaeological data for hand preference and great ape hand-use. The data show that skeletal asymmetries in arm and hand bones supporting a rightsided dominance were present at least in the genus Homo, although data are more robust for Neanderthals. The evidence from tool-use, production, and cave art confirms that right-hand preference was established in Neanderthals and was maintained until the present. The great apes can provide real-life models for testing the conditions that facilitate or enhance hand preference at both the individual and group levels. The database on great ape hand-use indicates that they do exhibit hand preferences, especially in complex tasks. However, their preferences vary between tasks, and while group-level biases have occasionally been reported, no human-like handedness bias has been found. We discuss the methodological problems encountered in these approaches. Shared problems include a lack of agreed terminology both within and between disciplines, small sample sizes, interpretation biases and a failure to replicate experiments. In general, there is a paucity of fossil material, with poor preservation hampering traditional metric methods. The archaeological data are often founded on unreliable methods. The primate database is plagued by the use of measures that could be inappropriate for revealing hand preference, and by methodological inconsistencies between studies. We emphasise the need to standardise the methods to allow between studies and species comparisons. We propose that when referring to "handedness" it is more appropriate to use the terms "hand preference" and "hand use", to avoid confusion with each discipline's own definition of handedness.

Research paper thumbnail of Hand Preferences for Bimanual Coordination in 77 Bonobos ( Pan paniscus ): Replication and Extension

International Journal of Primatology, 2011

The literature on manual laterality in nonhuman primates provides inconsistent and inconclusive f... more The literature on manual laterality in nonhuman primates provides inconsistent and inconclusive findings and is plagued by methodological issues (e.g., small samples, inconsistency in methods, inappropriate measures) and gaps. Few data are available on bonobos and these are only from small samples and for relatively simple tasks. We examined laterality in a large sample of bonobos for a complex task. We tested 48 bonobos from Lola Ya Bonobo sanctuary (DR Congo) in an extension of our previous study of 29 bonobos from 3 European zoos. We assessed hand preferences using the tube task, which involves bimanual coordination: one hand extracts food from a tube that is held by the other hand. This task is a good measure of laterality and it has been used in other studies. We recorded events (frequency) and independent bouts of food extraction. We found significant manual laterality, which was not influenced by the settings or rearing history. We observed little effect of sex and found an influence of age, with greater right hand use in adults. The laterality was marked, with strong preferences and most individuals being lateralized (when analyzing frequency). We found individual preferences, with no group-level bias, even when we combined the data from the sanctuary and the zoos to enlarge the sample to 77. These first data, for a complex task and based on a large sample, are consistent with previous findings in bonobos and in other nonhuman primate species for a variety of tasks. They suggest that, despite particular features in terms of proximity to humans, language and bipedalism, bonobos do not display a laterality that is more marked or more similar to human handedness compared to that of other nonhuman primate species.

Research paper thumbnail of Hand preferences for bimanual coordination in 29 bonobos ( Pan paniscus

Behavioural Brain Research, 2009

Brain lateralization has long been thought to be unique to humans. To investigate the origins and... more Brain lateralization has long been thought to be unique to humans. To investigate the origins and functions of this feature, researchers study behavioural laterality in other animals. Despite a substantial database, manual laterality in non-human primates remains a controversial topic. We give here a review of the main findings on manual preference in great apes. This article presents data on hand preferences for a bimanual coordination in 29 bonobos (Pan paniscus). The study aims to provide data on manual laterality for a complex bimanual task in this very interesting and rarely studied species. Hand preferences were assessed using the 'tube task'. This task has been used with other species, which allows reliable data comparisons. The task requires a bimanual coordinated precise action: the subject holds the tube with one hand while reaching for food inside with the other hand. As a complex task, this measure has been shown to be efficient in revealing hand preferences. It has revealed group-level right bias in chimpanzees. Bonobos had never been tested. We recorded both independent bouts (counting only the first pattern of a sequence of identical actions) and frequency (counting every action). The bonobos exhibited strong hand preferences. With frequency, 11 bonobos were classified as right-handed, 15 were left-handed and 3 had no preference. With bouts, 8 bonobos were right-handed, 9 were left-handed and 12 had no preference. No group-level bias appeared. The results are discussed in relation with previous findings and theories on brain lateralization.

Research paper thumbnail of Lateralization for visual processes: eye preference in Campbell’s monkeys ( Cercopithecus c. campbelli

Animal Cognition, 2009

Brain lateralization has been the matter of extensive research over the last centuries, but it re... more Brain lateralization has been the matter of extensive research over the last centuries, but it remains an unsolved issue. While hand preferences have been extensively studied, very few studies have investigated laterality of eye use in non-human primates. We examined eye preference in 14 Campbell's monkeys (Cercopithecus c. campbelli). We assessed eye preference to look at a seed placed inside a tube using monocular vision. Eye use was recorded for 100 independent and non-rewarded trials per individual. All of the 14 monkeys showed very strong preferences in the choice of the eye used to look inside the tube (mean preference: 97.6%). Eight subjects preferred the right eye and six subjects preferred the left eye. The results are discussed in light of previous data on eye preference in primates, and compared to data on hand preference from these subjects. Our findings would support the hypothesis for an early emergence of lateralization for perceptual processes compared to manual motor functions.

Research paper thumbnail of Manual laterality in Campbell's Monkeys ( Cercopithecus c. campbelli) in spontaneous and experimental actions

Behavioural Brain Research, 2006

Behavioural asymmetries, once thought to be exclusively human, appear to be widespread in vertebr... more Behavioural asymmetries, once thought to be exclusively human, appear to be widespread in vertebrates. A population-level bias should stem from natural selection and reflect a cerebral dominance, while individual preferences might be linked to individual characteristics. Manual laterality has been extensively investigated in non-human primates. However, despite a strong data base, no general patterns have emerged, resulting in a few explanatory theories and little consensus. This study was interested in manual laterality in 12 Campbell's monkeys (Cercopithecus c. campbelli). Several theories were examined, using both direct behavioural observations during feeding behaviour and six controlled experimental conditions, in which we varied task demands to investigate the effect of two factors. We systematically varied the individual posture (sat, tripedal, bipedal, clung) and the complexity of the task (box with or without a lid to open). Concerning the direction of preference, we found individual and action-specific preferences for experimental actions, which match previous reports. No population bias emerged and each subject appeared to react differently to the factors, hereby contradicting the theories. However, concerning the strength of preference, all individuals tended to be affected in similar ways. Spontaneous actions were less lateralized than experimental ones, and the simplest task and spontaneous category tended to show the weakest laterality. The relative complexity and novelty of these actions may account for these results.

Research paper thumbnail of Hand Preferences for Bimanual Coordination in 77 Bonobos (Pan paniscus): Replication and Extension

International Journal of Primatology, Jan 11, 2011

The literature on manual laterality in nonhuman primates provides inconsistent and inconclusive f... more The literature on manual laterality in nonhuman primates provides inconsistent and inconclusive findings and is plagued by methodological issues (e.g., small samples, inconsistency in methods, inappropriate measures) and gaps. Few data are available on bonobos and these are only from small samples and for relatively simple tasks. We examined laterality in a large sample of bonobos for a complex task. We tested 48 bonobos from Lola Ya Bonobo sanctuary (DR Congo) in an extension of our previous study of 29 bonobos from 3 European zoos. We assessed hand preferences using the tube task, which involves bimanual coordination: one hand extracts food from a tube that is held by the other hand. This task is a good measure of laterality and it has been used in other studies. We recorded events (frequency) and independent bouts of food extraction. We found significant manual laterality, which was not influenced by the settings or rearing history. We observed little effect of sex and found an influence of age, with greater right hand use in adults. The laterality was marked, with strong preferences and most individuals being lateralized (when analyzing frequency). We found individual preferences, with no group-level bias, even when we combined the data from the sanctuary and the zoos to enlarge the sample to 77. These first data, for a complex task and based on a large sample, are consistent with previous findings in

Research paper thumbnail of Hand preference and its flexibility according to the position of the object: a study in cercopithecines examining spontaneous behaviour and an experimental task (the Bishop QHP task)

The extant literature on manual laterality in non-human primates is inconclusive, plagued by inco... more The extant literature on manual laterality in non-human primates is inconclusive, plagued by inconsistent or contradictory findings and by disturbing methodological issues (e.g. uncontrolled influential factors, comparability issues). The present study examined hand preference and its flexibility in 15 red-capped mangabeys (C. t. torquatus) and 13 Campbell's monkeys (C. c. campbelli), two species that differ in their degree of arboreality. We investigated the influence of the spatial position of the object on hand preference for reaching. We considered spontaneous behaviour (reaching for food during daily feeding) and an experimental task: the QHP task. The QHP is a task that is used in humans. This is a simple reaching task that involves high spatial constraints on hand use. In our study, the subject had to reach for items that were placed on a semi-circle in front of it on five positions, including in the centre position, in the ipsilateral space and in the contralateral space. We assessed hand preference for reaching in front (baseline condition), and we examined how this preference changed when reaching in lateral positions. For reaching in front, about half of the subjects were lateralized and no group-level bias occurred, for both spontaneous and experimental conditions. When considering reaching in the lateral positions, we observed that the position of the object influenced hand use: individuals used the hand that was closest to the object. The results are discussed in relation to previous findings in humans and in non-human primates and regarding theories on handedness and flexibility of hand preference.

Research paper thumbnail of The evolution of handedness in humans and great apes: a review and current issues

Population-level right-handedness is a defining characteristic of humans. Despite extensive resea... more Population-level right-handedness is a defining characteristic of humans. Despite extensive research, we still do not know the conditions or timing of its emergence in human evolution. We present a review of research into the origins of handedness, based on fossil and archaeological data for hand preference and great ape hand-use. The data show that skeletal asymmetries in arm and hand bones supporting a rightsided dominance were present at least in the genus Homo, although data are more robust for Neanderthals. The evidence from tool-use, production, and cave art confirms that right-hand preference was established in Neanderthals and was maintained until the present. The great apes can provide real-life models for testing the conditions that facilitate or enhance hand preference at both the individual and group levels. The database on great ape hand-use indicates that they do exhibit hand preferences, especially in complex tasks. However, their preferences vary between tasks, and while group-level biases have occasionally been reported, no human-like handedness bias has been found. We discuss the methodological problems encountered in these approaches. Shared problems include a lack of agreed terminology both within and between disciplines, small sample sizes, interpretation biases and a failure to replicate experiments. In general, there is a paucity of fossil material, with poor preservation hampering traditional metric methods. The archaeological data are often founded on unreliable methods. The primate database is plagued by the use of measures that could be inappropriate for revealing hand preference, and by methodological inconsistencies between studies. We emphasise the need to standardise the methods to allow between studies and species comparisons. We propose that when referring to "handedness" it is more appropriate to use the terms "hand preference" and "hand use", to avoid confusion with each discipline's own definition of handedness.

Research paper thumbnail of Hand Preferences for Bimanual Coordination in 77 Bonobos ( Pan paniscus ): Replication and Extension

International Journal of Primatology, 2011

The literature on manual laterality in nonhuman primates provides inconsistent and inconclusive f... more The literature on manual laterality in nonhuman primates provides inconsistent and inconclusive findings and is plagued by methodological issues (e.g., small samples, inconsistency in methods, inappropriate measures) and gaps. Few data are available on bonobos and these are only from small samples and for relatively simple tasks. We examined laterality in a large sample of bonobos for a complex task. We tested 48 bonobos from Lola Ya Bonobo sanctuary (DR Congo) in an extension of our previous study of 29 bonobos from 3 European zoos. We assessed hand preferences using the tube task, which involves bimanual coordination: one hand extracts food from a tube that is held by the other hand. This task is a good measure of laterality and it has been used in other studies. We recorded events (frequency) and independent bouts of food extraction. We found significant manual laterality, which was not influenced by the settings or rearing history. We observed little effect of sex and found an influence of age, with greater right hand use in adults. The laterality was marked, with strong preferences and most individuals being lateralized (when analyzing frequency). We found individual preferences, with no group-level bias, even when we combined the data from the sanctuary and the zoos to enlarge the sample to 77. These first data, for a complex task and based on a large sample, are consistent with previous findings in bonobos and in other nonhuman primate species for a variety of tasks. They suggest that, despite particular features in terms of proximity to humans, language and bipedalism, bonobos do not display a laterality that is more marked or more similar to human handedness compared to that of other nonhuman primate species.

Research paper thumbnail of Hand preferences for bimanual coordination in 29 bonobos ( Pan paniscus

Behavioural Brain Research, 2009

Brain lateralization has long been thought to be unique to humans. To investigate the origins and... more Brain lateralization has long been thought to be unique to humans. To investigate the origins and functions of this feature, researchers study behavioural laterality in other animals. Despite a substantial database, manual laterality in non-human primates remains a controversial topic. We give here a review of the main findings on manual preference in great apes. This article presents data on hand preferences for a bimanual coordination in 29 bonobos (Pan paniscus). The study aims to provide data on manual laterality for a complex bimanual task in this very interesting and rarely studied species. Hand preferences were assessed using the 'tube task'. This task has been used with other species, which allows reliable data comparisons. The task requires a bimanual coordinated precise action: the subject holds the tube with one hand while reaching for food inside with the other hand. As a complex task, this measure has been shown to be efficient in revealing hand preferences. It has revealed group-level right bias in chimpanzees. Bonobos had never been tested. We recorded both independent bouts (counting only the first pattern of a sequence of identical actions) and frequency (counting every action). The bonobos exhibited strong hand preferences. With frequency, 11 bonobos were classified as right-handed, 15 were left-handed and 3 had no preference. With bouts, 8 bonobos were right-handed, 9 were left-handed and 12 had no preference. No group-level bias appeared. The results are discussed in relation with previous findings and theories on brain lateralization.

Research paper thumbnail of Lateralization for visual processes: eye preference in Campbell’s monkeys ( Cercopithecus c. campbelli

Animal Cognition, 2009

Brain lateralization has been the matter of extensive research over the last centuries, but it re... more Brain lateralization has been the matter of extensive research over the last centuries, but it remains an unsolved issue. While hand preferences have been extensively studied, very few studies have investigated laterality of eye use in non-human primates. We examined eye preference in 14 Campbell's monkeys (Cercopithecus c. campbelli). We assessed eye preference to look at a seed placed inside a tube using monocular vision. Eye use was recorded for 100 independent and non-rewarded trials per individual. All of the 14 monkeys showed very strong preferences in the choice of the eye used to look inside the tube (mean preference: 97.6%). Eight subjects preferred the right eye and six subjects preferred the left eye. The results are discussed in light of previous data on eye preference in primates, and compared to data on hand preference from these subjects. Our findings would support the hypothesis for an early emergence of lateralization for perceptual processes compared to manual motor functions.

Research paper thumbnail of Manual laterality in Campbell's Monkeys ( Cercopithecus c. campbelli) in spontaneous and experimental actions

Behavioural Brain Research, 2006

Behavioural asymmetries, once thought to be exclusively human, appear to be widespread in vertebr... more Behavioural asymmetries, once thought to be exclusively human, appear to be widespread in vertebrates. A population-level bias should stem from natural selection and reflect a cerebral dominance, while individual preferences might be linked to individual characteristics. Manual laterality has been extensively investigated in non-human primates. However, despite a strong data base, no general patterns have emerged, resulting in a few explanatory theories and little consensus. This study was interested in manual laterality in 12 Campbell's monkeys (Cercopithecus c. campbelli). Several theories were examined, using both direct behavioural observations during feeding behaviour and six controlled experimental conditions, in which we varied task demands to investigate the effect of two factors. We systematically varied the individual posture (sat, tripedal, bipedal, clung) and the complexity of the task (box with or without a lid to open). Concerning the direction of preference, we found individual and action-specific preferences for experimental actions, which match previous reports. No population bias emerged and each subject appeared to react differently to the factors, hereby contradicting the theories. However, concerning the strength of preference, all individuals tended to be affected in similar ways. Spontaneous actions were less lateralized than experimental ones, and the simplest task and spontaneous category tended to show the weakest laterality. The relative complexity and novelty of these actions may account for these results.