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Developmental Science 6:1 (2003), pp 55–61

REPORT
Blackwell Science, Ltd

From hand to mouth in the evolution of language: the influence

Laterality in gestural and vocal communication

of vocal behavior on lateralized hand use in manual gestures by

chimpanzees ( Pan troglodytes)
William D. Hopkins1,2 and Monica Cantero3
1. Division of Psychobiology, Yerkes National Primate Research Center, Atlanta, Georgia, USA
2. Department of Psychology, Berry College, Mount Berry, Georgia, USA
3. Department of Foreign Languages, Berry College, Mount Berry, Georgia, USA

Abstract
Studies in human subjects indicate that manual gestures accompanied by speech are produced more often by the right compared
to the left hand. Additional studies indicate that the production of sign language is controlled by the same brain areas as speech,
suggesting similar neurobiological substrates for language that are not modality specific. We report evidence that chimpanzees
exhibit preferential use of the right hand in gestural communication. Moreover, use of the right hand in gestural communication
is significantly enhanced when accompanied by a vocalization, particularly among human-reared chimpanzees. Taken together, the
data suggest that the lateralization of manual and speech systems of communication may date back as far as 5 million years ago.

Introduction that there are significantly more right- compared to left-

Clinical, experimental and developmental data all sug-
gest that motor systems involved in the control of man-
ual and oral movements are lateralized in the left
hemisphere (Kimura, 1993; Iverson & Thelen, 1999). In
humans, it has been well established that language com-
prehension and production are lateralized in distinct
regions within the left cerebral hemisphere (Corballis,
1992). Although initial clinical and experimental studies
focused on speech comprehension and production,
recent findings in deaf populations who have learned to
use non-speech communication systems suggest that
similar areas within the left hemisphere are involved in
comprehending and producing signs (Corina, Vaid &
Bellugi, 1992; Grossi, Semenza, Corazza & Volterra,
1996; Kimura, 1993). These results suggest that func-
tional asymmetries in communicative systems are not
modality specific and that similar neurobiological sys-
tems mediate manual and spoken languages.
Further evidence suggesting an association between
left hemisphere dominance in manual and spoken com-
munication comes from the reports of manual asym-
metries in mimetic gestures produced while subjects are
engaged in speech. For example, it has been reported
hand gestures while subjects verbally recall memory list
items or describe narrative text (Foundas, Macauley,
Raymer, Maher, Heilman & Rothi, 1995; Kimura, 1973).
These data have been interpreted as evidence that the pro-
duction of speech activates motor areas within the left
hemisphere that results in preferential use of the right
hand in simultaneous gesticulation. Finally, develop-
mental studies indicate that as infants grow older there
is a progression toward increased preferential use of the
right hand in gestural communication (Blake, O’Rourke
& Borzellino, 1994). Moreover, in infants, increased
right-hand use in gestures is associated with increased
use of speech or vocalizations (Dalby, Gibson, Grossi &
Schneider, 1980; Locke, Bekken, McMinn-Larson & Wein,
1995).
Whether homologous asymmetries are present in the
processing of communicative behavior by nonhumans
remains a topic of considerable debate, particularly as it
pertains to discussions of the neurobiology of language
origins (see Steklis & Raleigh, 1979a, 1979b; Bradshaw
& Rogers, 1993 for review). There are numerous reports
of left hemisphere specialization in the processing of
species-specific sounds as well as other classes of acoustic
stimuli (see Hopkins & Fernandez-Carriba, 2002 for

Address for correspondence: William D. Hopkins, Division of Psychobiology, Yerkes National Primate Research Center, Emory University, Atlanta,
Georgia, 30322, USA; e-mail: lrcbh@rmy.emory.edu or whopkins@berry.edu

© Blackwell Publishing Ltd. 2003, 9600 Garsington Road, Oxford OX 4 2DQ, UK and 350 Main Street, Malden, MA 02148, USA.
DESC_013.fm Page 56 Tuesday, January 28, 2003 11:46 AM
56 William D. Hopkins and Monica Cantero
review). Far fewer studies have examined asymmetries in
the production of communicative signals. Left hemi-
sphere asymmetries in the production of vocalizations
have been reported in some non-mammalian species
such as birds and frogs (Bisazza, Rogers & Vallortigara,
1998; Bradshaw & Rogers, 1993); however, in primates,
the evidence suggests right hemisphere asymmetries
in the production of vocalizations, particularly those
with negative emotional valence (Hauser, 1993; Hook-
Costigan & Rogers, 1998). These findings are consistent
with the general view that the primary function of prim-
ate vocalizations is in communicating affective rather
than semantic or other linguistic information (but see
Seyfarth & Cheney, 1997). In humans, the right hemi-
sphere plays a role in producing non-linguistic auditory
information while the left hemisphere is responsible
for speech and language production associated with
semantics and grammatical information.
In contrast to the functions of vocalizations in non-
human primates, recent studies indicate that great apes
use manual gestures to point to otherwise unattainable
things and they alternate their gaze between the referent
and communicative recipient (Call & Tomasello, 1994;
Krause & Fouts, 1997; Leavens & Hopkins, 1998). More
recent findings indicate that chimpanzees modify their
communicative behavior in accordance with the atten-
tional state of the audience such that vocal and other
auditory signals are used to capture the attention of an
inattentive recipient. In contrast, when the audience is
attentive to the communicator, chimpanzees and other
great apes will more frequently use manual gestures
rather than auditory signals to communicate to the
recipient (Hostetter, Cantero & Hopkins, 2001; Toma-
sello, Call, Nagell, Olguin & Carpenter, 1994; but see
Theall & Povinelli, 1999). Although more research is
needed to characterize the different factors that influ-
ence communicative behavior in great apes, some studies
indicate that chimpanzees and other great apes produce
manual gestures, and to a lesser extent vocalizations
and other acoustic signals, that may be both intentional
and referential, notable prerequisites for the evolution of
language and speech in humans. Based on these findings,
the potential for antecedents to the evolution of left
hemisphere lateralization in language and speech pro-
duction in humans may lie in the expression of prefer-
ential hand use for manual gestures in great apes, notably
chimpanzees.
The purpose of the current study was to further exam-
ine the association between hand use for gestural com-
munication and vocal behavior in chimpanzees. This was
prompted by observations that right-hand use in gestural
communication is found to be significantly higher when
the gestures are accompanied by a vocalization (Hopkins
© Blackwell Publishing Ltd. 2003
& Leavens, 1998). Unfortunately, in the Hopkins and
Leavens (1998) study, it is not clear whether the associ-
ation between right handed gesturing and vocal beha-
vior exists at a population level nor is it clear whether
the effect is due to the referential or affective expression
of the vocal behavior. Based on the existing literature
on lateralization in the production of vocalization in
nonhuman primates, two hypotheses can be proposed
about the relationship between vocal and gestural com-
munication in chimpanzees. Specifically, if the produc-
tion of vocalizations is mediated by the right hemisphere
(e.g. Hauser, 1993) then subjects that vocalize while ges-
turing should show a higher incidence of left- compared
to right-handed gesturing. In contrast, if the production
of vocalizations is mediated by the left hemisphere, then
right-handed manual gestures should be more prevalent
when subjects are simultaneously vocalizing compared
to when they are not. Lastly, hand use for gesture could
be independent of vocalization; therefore, no association
between hand use for gestures and vocalization should
be evident.
Methods
Subjects
The sample consisted of 73 chimpanzees housed at the
Yerkes National Primate Research Center (YNPRC)
ranging in age from 7 to 44 years. There were 38 females
and 35 males, respectively. Within the female sample there
were 13 human-reared and 25 mother-reared subjects. In
the male sample there were 23 human-reared and 12 mother-
reared chimpanzees. Mother-reared subjects were chim-
panzees that were raised by their biological, conspecific
mother beyond 30 days of life. Human-reared subjects
were chimpanzees that were either rejected by their bio-
logical mother or removed from their biological mother
for experimental purposes and raised by humans. Human-
reared chimpanzees are raised in standard nursery-
setting with other neonatal and infant chimpanzees until
3 years of age at which time they are integrated into multi-
age, multi-sex chimpanzee social groups (see Bard, 1996
for summary of rearing practices of human-raised chim-
panzees at the YNPRC).
Procedure
The testing procedure was relatively straightforward and
was similar to that used in previous studies on gestural
communication in this colony of chimpanzees (see Hop-
kins & Wesley, 2002; Leavens & Hopkins, 1998, 1999 for
reviews). On each test trial, an experimenter approached
DESC_013.fm Page 57 Tuesday, January 28, 2003 11:46 AM
the subject’s home cage and positioned themselves approx-
imately 1 m from the subject’s home cage and directly in
front of the focal chimpanzee. Every effort was made to
ensure that the experimenters were positioned centrally
in front of the subjects so as to preclude any effects of
positional bias on hand use. Moreover, the experimenter
approached the subject’s home cage from different direc-
tions across test trials to control for this potential extrane-
ous variable. After approaching, the experimenter offered
the chimpanzee a banana that was previously out of
sight from the chimpanzee. Prior to offering the banana,
the banana had been hidden inside the laboratory coat
pocket worn by the experimenter. Because the experi-
menter was approximately 1 m from the focal subject,
the banana was out of their immediate reach and this
condition stimulated the production of communicative
behaviors by the focal chimpanzee. The experimenter
held onto the banana throughout the entire duration of
the trial and did not move from the initial point from
which they offered the banana. The experimenter also
maintained eye contact with the subject throughout the
duration of the trial in order to increase the probability
of engaging the subjects to produce a communicative
behavior.
A second experimenter, standing directly behind the
experimenter offering the banana, recorded on a sheet of
paper all the communicative behavior of the focal subject
for a 1-minute sampling period. The communicative beha-
vior was sampled for 1 minute in order to increase the
probability that the chimpanzees would be observed to
gesture while either vocalizing or not. The second experi-
menter recorded all instances of either a gesture accom-
panied by a vocalization or a gesture not accompanied
by a vocalization. The second experimenter also made
note of the hand used to gesture by the subject. A gesture
accompanied by a vocalization was only recorded when
the two events occurred simultaneously or within a 2-sec
interval. Moreover, only a single response was recorded
although, in principle, more than one vocalization could
be produced by a single gestural event. For example, if the
focal chimpanzee gestured and then produced multiple
vocalizations in a sequential manner, then only a single
gesture + vocal event was recorded. However, if the chim-
panzee stopped gesturing by pulling their hand away from
the cage mesh and then initiated another gesture, another
event was recorded. No attempt was made to characterize
the vocalizations in terms of their structure using spec-
trographic analysis. However, none of the vocalizations
would be considered species-specific such as food calls or
grunts and not all of the subjects were observed to vocalize.
Rather, among the subjects that did vocalize, the calls
varied from individual to individual and appeared to be
idiosyncratic sounds acquired in response to their captive
© Blackwell Publishing Ltd. 2003
Laterality in gestural and vocal communication 57
rearing. At the end of each test trial, the focal subject
was given the banana no matter what behaviors they
exhibited during the 1-min test session.
Each chimpanzee was tested on 10 separate occasions
separated by at least one day. The 10 test sessions were
administered over a 6-month period between July and
December 2000. Individual subjects were tested in their
home cages. Nearly all of the subjects lived in social
groups ranging from 2 to 16 individuals. Thus, focal
sampling techniques were used with one subject from
each cage serving as a subject on a given trial. Inter-rater
reliability in coding the hand use and frequency of
vocalizations was established in 10% of the sample of
subjects. Cohen’s kappa values were 0.994 and 0.850
when coding for hand use and the number of vocaliza-
tions produced. Both of these values are considered high
and indicative of good reliability.
Data analysis
The frequency data were summed across the 10 test sessions
and several different means of characterizing the data
were used in this study. First, general descriptive ana-
lyses were performed on the overall communicative beha-
viors, irrespective of hand use. Three variables were
derived including the total number of gestures (Total),
total number of gestures accompanied by a vocalization
(Sum + Vocal) and the total number of gestures produced
without a vocalization (Sum + No-Vocal). In terms of
laterality, handedness indices were derived by subtract-
ing the number of left-hand responses from the number
of right-hand responses and dividing by the total number
of responses HI = [(R − L)/(R + L)]. A total handedness
index was derived for all gestures (Sum-HI). In addition,
handedness indices were derived for gestures that were
accompanied by a vocalization (Gesture + Vocal) and
for those that were not (Gesture + No-Vocal). All ana-
lyses were done using parametric statistics with alpha set
to p < 0.05. Post-hoc analyses were performed using
Tukey’s HSD ( p < 0.05) unless otherwise noted.
Results
Descriptive statistics
The chimpanzees produced a total of 2,468 gestures
including 1,934 that were not accompanied by a vocaliza-
tion and 532 that were accompanied by a vocalization.
The mean number of gestures produced with and without
a vocalization as a function of sex and rearing history
can be seen in Table 1. Overall, there were no significant
sex or rearing differences in the total number of gestures
DESC_013.fm Page 58 Tuesday, January 28, 2003 11:46 AM
58 William D. Hopkins and Monica Cantero
Table 1 Mean number of gestures produced with and without
a vocalization
Variable Sum + Vocal Sum + No-Vocal
Sex
Males 8.01 24.09
Females 6.30 29.55
Rearing history
Human-Reared 3.22 28.24
Mother-Reared 11.09 25.40
produced by the chimpanzees. The total number of gestures
produced with and without a vocalization was also com-
pared using a mixed design analysis of variance. Sex and
rearing history served as between-group variables while
Sum + Vocal and Sum + No-Vocal served as repeated
measures. This analysis revealed a significant interaction
between communicative response and rearing history
F(1, 69) = 4.79, p < 0.01. Post-hoc analysis indicated that
mother-reared chimpanzees produced more Sum + Vocal
responses compared to human-reared subjects. No sig-
nificant differences between mother- and human-reared
chimpanzees were found for the Sum + No-Vocal measure.
In short, mother-reared produced more gestures that
were accompanied by a vocalization than human-reared
chimpanzees.
Laterality effects
One-sample t-tests were performed on the handedness
scores to assess population-level biases in hand use.
For the Sum-HI t(72) = 3.92, p < 0.001, Gesture + Vocal
t(50) = 4.53, p < 0.001 and Gesture + No-Vocal t(72) =
2.92, p < 0.01 measures, significant right-hand popula-
tion biases were found. In the next analysis, whether the
magnitude in right-hand use differed within subjects that
gestured while vocalizing or not was assessed. Within the
entire sample of chimpanzees, 51 subjects produced ges-
tures both with and without vocalizations. This allowed
for a within-subject comparison of the influence of vocal
behavior on the expression of hand preference in gestural
communication. For this analysis, the Gesture + Vocal and
Gesture + No-Vocal handedness scores were compared
using a paired samples t-test. The analysis revealed a
significant difference with the Gesture + Vocal handed-
ness scores significantly higher than the Gesture +
No-Vocal handedness scores t(50) = 3.51, p < 0.01 (see
Figure 1).
Sex and rearing effects of lateralized hand use
For this analysis, the handedness index values for the
responses accompanied by (Gesture + Vocal) or not
© Blackwell Publishing Ltd. 2003
Figure 1 The mean HI values for the overall number of
gestures (Sum-HI), number of gestures produced with a
vocalization (Gesture + Vocal) and the number of gestures
produced without a vocalization (Gesture + No-Vocal). All
were significantly different from zero, even after correcting for
Type I error using Bonferonni’s correction procedure. In
addition, the HI values for the Gesture + Vocal responses were
significantly higher than the HI values produced for the
Gesture − Vocal responses, as revealed by a paired samples
t-test.
accompanied by a vocalization (Gesture + No-Vocal)
were analyzed using two separate 2 (sex) × 2 (rearing
history) analyses of covariance (ANCOVA). The cov-
ariate was the Sum + Vocal and Sum + No-Vocal values
from the initial analysis. ANCOVA was used to statist-
ically control for differences in the total number of
responses produced by mother- and human-reared chim-
panzees and its potential influence on the handedness
index values. For the Gesture + Vocal data, significant
main effects for sex F(1, 46) = 5.58, p < 0.03 and rearing
history F(1, 46) = 6.23, p < 0.02 were found. Females
(Mean = 0.71) had higher HI values then males (Mean =
0.21) and human-reared chimpanzees (Mean = 0.74)
had higher HI values than mother-reared chimpanzees
(Mean = 0.19). For the Gesture + No-Vocal responses,
no significant main effects for rearing history or sex were
found.
Comparisons with other measures of hand use
The results of the previous analyses on the Gesture +
Vocal and Gesture + No-Vocal measures indicated that
human-reared chimpanzees were more right-handed
than mother-reared individuals. What is not clear is
whether this effect generalizes to other motor tasks or is
specific to gestural communication. Specifically, it could
be argued that these effects are due to the fact that the
chimpanzees are being reared in a human designed,
right-handed world (see Collins, 1985). If this explana-
tion accounts for the observed findings then the rearing
DESC_013.fm Page 59 Tuesday, January 28, 2003 11:46 AM
differences should be evident for measures of hand use
that are not communicative in function. In contrast, if
the rearing effect is specific to gestures accompanied by
a vocalization then the importance of human rearing on
the development of laterality would be limited to the
expression of communicative behavior.
To test this hypothesis, the handedness scores for
human- and mother-reared chimpanzees were com-
pared for three other measures of hand preference includ-
ing simple reaching (REACH), coordinated bimanual
actions (TUBE) and bimanual feeding (FEED). The
behavioral tasks used to assess hand preferences in the
chimpanzees have been described in detail elsewhere
(Hopkins, 1993, 1994, 1995a, 1995b). Listed below is a
brief description of each task and the general procedure
used to assess hand preference.
Bimanual feeding (FEED)
Each afternoon, the primates housed at the YRPRC
receive fruits and vegetables as part of their daily diet.
Each subject usually receives 2 oranges, 1 banana, some
celery stalks and/or carrots. Once retrieving the food, the
subjects typically move to a seating place and consume
the food. The chimpanzees typically hold the extra
pieces of food with one hand and feed with the opposite
hand. Hand use was recorded when the subjects were
feeding with one hand for a minimum duration of 3 seconds
and the nonfeeding hand was holding the remaining
portions of food. The dominant hand was recorded as
the one feeding.
Simple reaching (REACH)
For this measure, a piece of food (a peanut or raisin) was
randomly thrown into the subject’s cage. The subjects
had to locomote to the location and pick up the food.
The hand used to pick up the food was recorded. A trial
was counted only when the subjects maintained a tri-
pedal posture while reaching (i.e. one upper limb and
two hind limbs were on the floor).
Coordinated bimanual tube task (TUBE)
Poly-vinyl-chloride tubes (24 to 31 cm long, 2.5 cm wide)
with peanut butter smeared on the inside were given to
the subjects in their home cage. The digit and hand used
to remove the peanut butter was recorded each time the
subjects inserted their finger, removed peanut butter
from the tube and placed their finger in their mouth.
Observations continued until the subjects stop showing
interest in the tube (usually when they have eaten all the
peanut butter), dropped it for at least 10 seconds, or
© Blackwell Publishing Ltd. 2003
Laterality in gestural and vocal communication 59
pushed the tube back out of their home cage through
the cage mesh.
For all measures, a minimum of 25 responses was
collected in each subject. As with the gesture data, hand-
edness indices were derived following the procedure
described above. Positive values reflected right-hand
biases and negative values reflected left-hand biases. The
absolute value reflected the strength in hand preference.
Separate 2 (sex) × 2 (rearing history) ANOVAs for each
handedness measure failed to reveal any significant
main effects or interactions. Thus, the effect of human
rearing on lateralized hand use is specific to gestural
communication.
Discussion
The findings from this study indicate that preferential
use of the right hand for gestures is significantly enhanced
when the gestures are accompanied by a vocalization,
particularly among chimpanzees raised by humans. This
effect cannot be attributed to an overall influence of
human rearing on hand preference because differences
in handedness were not found for other measures of
hand use. Moreover, hand use for gestures did not posit-
ively correlate with any other measures of hand use.
Based on previous studies examining the association
between vocal production and manual gesticulation in
humans, the evidence of increased right-hand use in ges-
tures that are accompanied by a vocalization suggest
that the left hemisphere is controlling both the manual
and vocal communication systems by chimpanzees. If
there was bilateral or right hemisphere control of the
vocalizations then right-hand gestures should have been
inhibited or reduced in the chimpanzees. We believe this
is the first evidence of asymmetries in the production of
intentional, referential communicative signals by nonhu-
man primates.
It could be argued that the differences in handedness
scores between mother- and human-reared chimpanzees
are due to a sampling bias in total hand use because the
mother-reared chimpanzees produced significantly more
gestures accompanied by a vocalization than human-
reared chimpanzees. Although this explanation cannot
be ruled out, we believe it is unlikely for three reasons.
First, the total number of overall responses does not
correlate with the handedness scores derived for gestures
produced with (r = −0.026, df = 49, n.s.) or without a
vocalization (r = 0.087, df = 71, n.s.). If sampling bias
explained the results then positive correlations should
have been evident. Second, the ANCOVA did not reveal
a significant influence of the covariate (total number of
responses) on the handedness scores. Third, the mean
DESC_013.fm Page 60 Tuesday, January 28, 2003 11:46 AM
60 William D. Hopkins and Monica Cantero
handedness scores for gestures produced with (Mean =
0.460; Adjusted Mean = 0.461) and without a vocaliza-
tion (Mean = 0.186; Adjusted Mean = 0.185) were nearly
identical whether or not the total number of responses
was used to adjust the handedness scores.
What is unclear from this study is why the human-
reared chimpanzees were more right handed than the
mother-reared chimpanzees when gestures were accom-
panied by a vocalization. One explanation may be that
the human-reared chimpanzees use their vocalizations in
conjunction with their gestures for different functions
compared to mother-reared chimpanzees. In some previ-
ous research with many of the same chimpanzees as those
used in this study, it has been found that human-reared
chimpanzees were more likely to exhibit gaze alternation
between a referent and human experimenter than mother-
reared chimpanzees (Leavens & Hopkins, 1998). It may be
that the human-reared chimpanzees are using their vocal-
izations in a functionally distinct way that has an influence
on their hand use. Specifically, the human-reared chimpan-
zees may be attempting to use their vocalizations in a
referential manner while the mother-reared chimpanzees
are using their vocalizations to communicate affective.
The referential use of the vocalizations by the human-
reared chimpanzees may induce left hemisphere activa-
tion, which results in greater use of the right hand for
gestures accompanying these calls. No attempt was
made to characterize the type of vocalizations made by
the chimpanzees nor was any attempt made to code for
the affective valence of the vocalizations. Future studies
should focus on this potential relationship between the
functional use of vocalizations and hand use.
In conclusion, the results indicate that preferential use
of the right hand is enhanced when accompanied by a
vocalization in chimpanzees, particularly in those raised by
human subjects. The preferential use of the right hand
for gestural communication did not correlate with other
measures of hand use and therefore does not appear to
reflect a bias associated with other motor functions (also
see Hopkins & Wesley, 2002). This indirectly supports
the theoretical notion that lateralization for referential
communication may have a unique neurobiological rep-
resentation in the brain compared to other behavioral,
motor and emotional functions. What brain areas may
be associated with asymmetries in communication is not
clear. With the recent evidence of left hemisphere neuro-
anatomical asymmetries in the peri-sylvian fissure region
as well as in the inferior frontal lobe, these appear to
be ideal regions for future consideration (Cantalupo &
Hopkins, 2001; Gannon, Holloway, Broadfield & Braun,
1998; Hopkins, Marino, Rilling & MacGregor, 1998;
Hopkins, Pilcher & MacGregor, 2001; Yeni-Komshian
& Benson, 1976).
© Blackwell Publishing Ltd. 2003
Acknowledgements
This research was supported by NIH grants NS-29574,
NS-36605 and RR-00165 to the Yerkes Regional Primate
Research Center. We thank Michael J. Wesley and
Autumn Hostetter for assistance in data collection. The
Yerkes Center is fully accredited by the American Asso-
ciation for Accreditation of Laboratory Animal Care.
APA guidelines for the ethical treatment of animals were
adhered to during all aspects of this study.
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Received: 12 April 2001
Accepted: 19 December 2001