Behavioural Processes 91 (2012) 202–205

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Behavioural Processes
journal homepage: www.elsevier.com/locate/behavproc

Short report

Hemispheric specialization in domestic dogs (Canis familiaris) for processing

different types of acoustic stimuli
Anna Reinholz-Trojan a , Ewelina Włodarczyk b , Maciej Trojan a , Adam Kulczyński a , Joanna Stefańska c,∗
a
University of Warsaw, Faculty of Psychology, ul. Stawki 5/7, 00-183 Warszawa, Poland
b
Interdisciplinary Center of Ethology and Animal Psychology, Aleja Stanow Zjednoczonych 24, 01-001 Warszawa, Poland
c
ALTO Center for Dog Education, ul. Nowoursynowska 115A, 02-797 Warszawa, Poland1

a r t i c l e i n f o a b s t r a c t

Article history: Lateralization is considered to be a fundamental feature of vertebrate brains. The aim of the present
Received 29 January 2012 study was to examine the impact of functional cerebral asymmetry on processing of auditory stimuli in
Received in revised form 2 July 2012 the domestic dog (Canis familiaris) during the orientation reaction. The experiment was conducted on
Accepted 3 July 2012
46 dogs (25 females and 21 males). Four types of auditory stimuli were used in the experiment (three
meaningful stimuli: cat meowing, dog barking, the “sit” command (“siad” in Polish), and a neutral word
Keywords:
(“wir”, meaning “whirl” in Polish). It was predicted that the orientation reaction (turning the head towards
Dog behaviour
the stimuli) would take place only in the case of meaningful sounds. It was also expected that dogs would
Functional asymmetry
Lateralization
show consistent lateralization. As predicted, all three meaningful stimuli elicited the orientation reaction.
Orientation reaction The response of the examined dogs to cat meowing showed significant lateralization with dominant
Reaction to auditory stimuli leftwards movement, which hints towards activation of the right cerebral hemisphere and may be related
Reaction to sound to strong emotions evoked by this stimulus. Contrary to results of previous studies, dogs reacting to
dog barking turned their heads leftwards more often, which suggests activation of the right cerebral
hemisphere, probably related to the emotional meaning of the stimulus. The “sit” command consistently
evoked the orientation reaction but there was no significant lateralization of this movement.
© 2012 Elsevier B.V. All rights reserved.

1. Introduction
Lateralization, also known as functional cerebral asymmetry, is
defined as the existence of differences in the functioning of the
brain hemispheres that are manifested in observable differences in
the use of organs located on each side of the body (hand, eye, ear,
paw, etc.). Diagnosing cerebral lateralization based on side pref-
erence in observed behaviours has been used in experiments and
tests on humans for over 50 years (e.g., Zazzo et al., 1960) and on
animals for nearly 20 years (e.g., Basile et al., 2009; Böye et al., 2005;
Branson and Rogers, 2006; Hauser and Anderson, 1994; Palleroni
and Hauser, 2003). Lateralization of brain functions and related
behaviours is present in many animal species and is a fundamen-
tal feature of vertebrate brains that enables parallel processing of
information, efficient classification of novel versus familiar objects
and initiating appropriate reactions (MacNeilage et al., 2009).
However, relatively little is known about lateralization in
domestic dogs. The study of Wells (2003) confirmed that the effect
of lateralization was observed for most examined dogs but there
was no clear preference for a particular paw at the population level.
∗ Corresponding author. Tel.: +48 607234104.
E-mail address: jstefanska@psych.uw.edu.pl (J. Stefańska).
1
Home address.
Other studies report that preference for using the right paw was
recorded in dogs (Tan, 1987).
One of the possible explanations for the observed differences
(domination of left or right hemisphere) may be related to sex dif-
ferences. Recent findings suggest that there is a tendency to use the
right paw in female rats (Elalmis et al., 2003) and horses (Murphy
et al., 2005). Both Wells (2003) and Quaranta et al. (2004) reported
that male dogs tended to prefer using the left paw and females
the right one. These differences may be caused by the effect of
hormones on brain functioning (Rogers and Andrew, 2002).
The present study is focused on the lateralization of processes
related to detection of vocalizations produced by specimens of the
same species and different species. Numerous studies reported
dominance of the left hemisphere in reception of vocalizations
produced by same-species specimens of birds, sea-lions, and mice
(Böye et al., 2005; Ehret, 1987; George et al., 2002; Palleroni
and Hauser, 2003). Activation of the left hemisphere has been
detected in primates generating some types of social communi-
cations. Hook-Costigan and Rogers (1998) noticed that Catarrhines
opened the right side of their mouth wider than the left one when
calling other monkeys. The reverse pattern was observed during
calls expressing fear.
Hauser and Anderson (1994) studied the orientation reaction
in rhesus monkeys. This reaction is inborn elementary behaviour
that involves directing telesensors towards a novel stimulus.

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 A. Reinholz-Trojan et al. / Behavioural Processes 91 (2012) 202–205
Researchers reported that the left hemisphere of these monkeys
was more active, as indicated by higher frequency of turning the
head rightwards in response to various calls emitted by other rhe-
sus monkeys (including aggression, fear, and orientation calls).
Alarm calls of a marine bird (Arenaria intepres) living in the same
environment as the rhesus monkeys generated leftwards head
turns more often (which indicated activation of the right hemi-
sphere). Psychophysiological experiments on Japanese macaques
(Macaca fuscata) also confirmed domination of the left hemisphere
in processing vocalization of animals of the same species (Hauser
et al., 1998).
Experiments thus indicate that activation of the right hemi-
sphere can be related to responding to unexpected, novel stimuli
and expression of strong emotions, such as aggression or fear. The
left hemisphere is activated for processing invariant, familiar and
repeated stimuli.
These results inspired Siniscalchi et al. (2008), who examined
whether dogs used different hemispheres for processing auditory
stimuli such as thunderstorm and barking of other dogs. The exper-
iment was performed on 14 dogs (8 females and 6 males) that were
exposed to recordings of three different canine vocalizations: bark-
ing in response to a stranger knocking on the door, dog vocalizing
when isolated, and dog vocalizing during play. The fourth record-
ing was the sound of thunder. The examination was performed at
the location where the dogs lived, in yards. The results were con-
sistent with predictions – dogs turned their heads to the right in
response to other dogs’ vocalizations and to the left in response to
the thunderstorm sound.
This result is consistent with experiments on other species and
it probably means that activation of the right hemisphere is related
to reactions to novel stimuli and expression of “intensive” emo-
tions, such as aggression, fear, and escape behaviour. There was no
significant relationship between paw preference in the examined
dogs (as tested beforehand) and lateralization of sound processing.
Our study aimed to examine how the orientation reaction
depends on the type of stimulus, in particular in terms of later-
alization. The experiment was performed in a laboratory setting.
We decided to test a larger sample of dogs than Siniscalchi et al.
(2008) and we used different stimuli. However, after a pilot study
we decided to abandon the multiple presentation of the stimuli to
avoid habituation. The results of Siniscalchi et al. (2008) convinced
us to abandon the initial test of paw preference in order to shorten
the experimental procedure and the time the animals had to spend
in the laboratory.
We expected that the orientation reaction would only be
observed in cases of significant stimuli – those that carried some
meaning or were emotionally exciting. These included barking,
meowing, and the “sit” command. Consistently with previous
results, we expected that dog barking would lead to a preference
for turning the head rightwards (common intraspecies social sig-
nal), and cat meowing, leftwards (emotionally stimulating sound).
In the case of the “sit” command, we expected that it would be pro-
cessed mainly by the left hemisphere, so that dogs would rather
turn their heads to the right, treating the command as an inter-
species communication. In the case of the neutral word “wir”, we
expected no lateralized orientation reaction. Consistently with the
results of Siniscalchi et al. (2008) we expected no sex differences.
2. Method
2.1. Animals
The experiment was performed on 46 domestic dogs of differ-
ent breeds (including mongrels), aged 2–8 years, including 25 intact
203
females and 21 intact males. The dogs were brought to the labora-
tory by their owners, who were also present during the experiment.
2.2. Apparatus and stimuli
The experiment was performed in an adapted room with the
following arrangement. The central spot in the room was occupied
by an agility table (90 cm × 90 cm, 40 cm high) on which the dogs
were placed. One metre behind the head of the dog standing on the
table was a wireless loudspeaker. The video-camera that recorded
animals’ behaviour was placed behind the loudspeaker (similarly
to the setup in Hauser and Anderson, 1994). The experimenter’s
table and the computer were located on the opposite side of the
room.
Dogs in our experiment heard a barking dog, a meowing cat, and
two words of human language; all sounds came from animals or
humans unfamiliar to the dogs tested in the experiment. The bark
was a recorded response of a dog to the appearance of a strange
dog. The cat meowing was the kind of threatening sound cats make
when approached by unwelcome intruders. The “siad” (“sit” in Pol-
ish; rhymes with English “bat”) command and the neutral word
“wir” (meaning “whirl” in Polish; rhymes with English/French “kir”)
were spoken by a male voice. The latter word is clearly different
from typical commands with which dogs might be familiar and we
assumed that it was meaningless to the examined dogs. The word
“wir” was therefore the control stimulus. All dogs were exposed to
the same recordings but the order of stimuli was randomized for
each dog.
2.3. Procedure
The research assistant led the owner and the dog to the experi-
mental room. Each dog was given about 10 min to become familiar
with the room. The dog was placed on the agility table and the
owner was asked to sit in front of it and to command the dog to
“stay” so that the animal would remain motionless in a position
parallel to the room axis. The owner was requested to focus the
dog using commands and/or treats so that the dog’s head would be
aligned exactly in the desired direction. When the dog was still, one
stimulus of volume 60 dB, randomly selected from the pool of four
experimental stimuli, was emitted. After the sound was emitted,
the dog was allowed to get off the table and play with the owner
for a few minutes. This procedure was repeated for each of the four
stimuli and each dog heard each stimulus only once. The reaction
of turning the head towards the loudspeaker was recorded, as well
as the direction of head rotation.
3. Results
The hypothesis linking the type of stimulus with the occur-
rence or absence of reaction was tested using the Q Cochran test,
assuming the occurrence of orientation reaction is coded as success.
The test result, Q(3) = 49, p = .001, indicates significant differences
between reactions to different stimuli (Fig. 1). Pairwise compar-
isons reveal that dogs reacted to the word “wir” (whirl) significantly
less frequently than all other stimuli. There were no significant dif-
ferences between reactions to other stimuli – the “sit” command,
dog barking, and cat meowing.
The preference for direction of head rotation was tested using
a single sample 2 test for all observations in which an orienta-
tion reaction was observed in response to the particular stimulus.
For the word “wir”, the result, 2(1) = 1.800, p = .180, suggests that
there is no preference for the direction of head rotation and ori-
entation of telesensors in the tested animals. Similar results were
obtained for the “sit” command (2(1) = 2.778, p = .096). However,
204 A. Reinholz-Trojan et al. / Behavioural Processes 91 (2012) 202–205
Fig. 1. Frequency of orientation reaction as a response to the four examined stimuli.
The dogs heard each stimulus only once and the order of stimuli was randomized
for each dog. More than half of the dogs ignored the word “whirl”. The frequency of
reactions to all other stimuli was significantly higher.
cat meowing elicited head rotation to the left more frequently than
to the right (2(1) = 5.565, p = .018). Lateralization was also observed
in reactions to dog barking (2(1) = 4.667, p = .031). Contrary to our
expectations, dogs turned their heads leftwards more frequently
than rightwards (Fig. 2).
No sex differences were detected for either the frequency of
orientation reaction or for the preference for head rotation.
4. Discussion
The present study aimed to examine whether dogs reacted to
calls produced by another dog and representatives of other species
(cat and human) and whether these reactions were lateralized. The
results indicate that the orientation reaction is observed signifi-
cantly more frequently in response to stimuli that are meaningful
for the recipient. A stimulus that has no valence and no semantic
meaning can be ignored, as demonstrated by comparing the neu-
tral word “wir” to other stimuli used in the experiment, which were
meaningful to the dogs.
It was expected that animals would turn their heads rightwards
in response to the “sit” command due to activation of the left
hemisphere, which in most species is responsible for processing
Fig. 2. Direction of head rotation in the orientation reaction for different types of
stimuli. Reactions to dog barking and cat meowing were lateralized – dogs turned
leftwards more frequently than rightwards. Reactions to human words showed no
side preference. No sex differences were detected.
familiar stimuli (Vallortigara et al., 2008). This effect was not con-
firmed in the present study (the result was not significant). Some
studies (e.g., Adams et al., 1987) focus on the acoustic character-
istics of human speech rather than the semantics and their results
indicate that the acoustic parameters of human vocalization are
processed in the right hemisphere of the dog brain. Thus, perhaps
it is the interference of the semantic meaning of the command with
the acoustic characteristics of the stimulus that is responsible for
the non-significant result in our study. Simultaneous activation of
the right hemisphere could be related to the novelty of the voice,
while the left hemisphere activation was due to familiar content of
the command. The expected result for cat meowing was that dogs
would tend to turn their heads leftwards, indicating activation of
the right hemisphere. Our results confirmed this hypothesis. It is
noteworthy that this stimulus invoked an orientation reaction in all
examined dogs. Previous studies indicate that the right hemisphere
is responsible for processing familiar interspecies calls (Hauser and
Anderson, 1994), and also, more importantly, for processing emo-
tionally significant stimuli (Hauser, 1993; MacNeilage et al., 2009;
Quaranta et al., 2007). Indeed, it can be reasonably assumed that an
angry meowing cat could be both a familiar and a significant type
of stimulus for dogs.
The predicted response to dog barking was that the dog would
turn its head rightwards, as observed in previous studies (Böye
et al., 2005; Ehret, 1987; George et al., 2002; Hauser and Anderson,
1994; Palleroni and Hauser, 2003; Petersen et al., 1978; Poremba
et al., 2004). However, we obtained the opposite result – dogs
turned their heads leftwards, indicating activation of the right
hemisphere. This result may be explained by the choice of stim-
ulus – perhaps the vocalization used in our study had emotional
significance to the dogs, evoking fear or aggression. This interpre-
tation is supported by the reaction of one dog participating in the
study, who tried to escape from the experimental table after being
exposed to the dog barking stimulus. Another possible explanation
relates to the fact that dogs participating in the study could have
been generally emotionally aroused due to the novel and artificial
circumstances of the testing environment and this arousal affected
their responses.
The present experiment provides evidence of functional asym-
metry in processing of auditory stimuli significant for the animal.
However, details of this process in the domestic dog (Canis famil-
iaris) seem to remain understudied and deserve further research.
In particular, certain aspects of the present study could be further
tested and improved – the results could be more reliable if the pro-
cedure were repeated several times on each animal. On the other
hand, multiple expositions to the stimulus might lead to habitu-
ation, which would extinguish elementary reactions such as the
orientation reflex.
Also, the selection of experimental stimuli could be further
improved to avoid emotional reactions to vocalizations that should
not elicit them. An interesting extension of this study could exam-
ine various aspects of different types of barking that may affect
lateralization of the response – such as emotional value, level of
arousal of the barking dog, meaning (type of barking). Perhaps
vocalizations could be selected by expert raters based on recordings
of vocalizations together with video recordings of the situations
where they were recorded. Further studies are also required regard-
ing reactions to spoken words.
Acknowledgments
The authors would like to thank the two anonymous review-
ers for their valuable comments and suggestions to improve the
manuscript. This study was funded by the Faculty of Psychology,
University of Warsaw, BST grant number 2010/524.
 A. Reinholz-Trojan et al. / Behavioural Processes 91 (2012) 202–205
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