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Relationship Between Paw Preference Strength and Noise Phobia in Canis familiaris


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The authors investigated the relationship between degree of lateralization and noise phobia in 48 domestic dogs (Canis familiaris) by scoring paw preference to hold a food object and relating it to reactivity to the sounds of thunderstorms and fireworks, measured by playback and a questionnaire. The dogs without a significant paw preference were significantly more reactive to the sounds than the dogs with either a left-paw or right-paw preference. Intense reactivity, therefore, is associated with a weaker strength of cerebral lateralization. The authors note the similarity between their finding and the weaker hand preferences shown in humans suffering extreme levels of anxiety and suggest neural mechanisms that may be involved.
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Relationship Between Paw Preference Strength and
Noise Phobia in Canis familiaris
N. J. Branson and L. J. Rogers
University of New England, Armidale, Australia
The authors investigated the relationship between degree of lateralization and noise phobia in 48
domestic dogs (Canis familiaris) by scoring paw preference to hold a food object and relating it to
reactivity to the sounds of thunderstorms and fireworks, measured by playback and a questionnaire. The
dogs without a significant paw preference were significantly more reactive to the sounds than the dogs
with either a left-paw or right-paw preference. Intense reactivity, therefore, is associated with a weaker
strength of cerebral lateralization. The authors note the similarity between their finding and the weaker
hand preferences shown in humans suffering extreme levels of anxiety and suggest neural mechanisms
that may be involved.
Keywords: noise phobia, lateralization, paw preference, dog, fear
Functional asymmetries of the cerebral hemispheres have been
identified in a number of vertebrate species using various methods,
including brain lesions, monocular testing, and turning biases
(summarized by Rogers & Andrew, 2002). Observations of hand,
or paw, preference have also been used as an indicator of hemi-
spheric asymmetry, particularly in primates (Hopkins & Bennett,
1994) but also in lower vertebrates (Bisazza, Cantalupo, Robins,
Rogers, & Vallortigara, 1997). Preferred use of one hand, or paw,
is associated with greater activity of the contralateral motor cortex
and is presumed to recruit activity in other regions of that hemi-
sphere (Yousry et al., 1995). As demonstrated in primates, use of
one hand induces changes in the size and distribution of movement
representations in the contralateral motor cortex (Nudo, Jenkins,
Merzenich, Prejean, & Grenda, 1992). We know, in humans, that
repeated movement of the fingers of one hand increases the blood
flow to the contralateral hemisphere (Halsey, Blauenstein, Wilson,
& Wills, 1979), which reflects greater neural activity in that
hemisphere. Papousek and Schulter (1999) showed that the
strength of right-handedness in humans is related to patterns of
asymmetrical hemispheric activity, measured by electroencepha-
logram. Using functional magnetic resonance imaging in humans,
it has been shown that strongly lateralized subjects exhibit cortical
activation of the primary and supplementary motor area contralat-
eral to the hand being moved to grasp an object, whereas the
ipsilateral hemisphere was more active in ambilateral subjects
(Nakada, Fujii, & Kwee, 2004).
Hand preference, in animals, is associated with temperament
and the type of behavior expressed in novel contexts: Marmosets
(Cameron & Rogers, 1999) and chimpanzees (Hopkins & Bennett,
1994) with a left (L)-hand preference to pick up and hold food
have been shown to interact with novel objects less readily than
those with a right (R)-hand preference. Moreover, positive corre-
lations have been shown between left-handedness and levels of
submissive behavior and the incidence of receiving aggression in
male rhesus macaques (Westergaard et al., 2003), although we
note that the same authors identified the opposite relationship in
female rhesus macaques (Westergaard et al., 2004). A relationship
has also been found between the use of one hand and the expres-
sion of emotion in humans: Schiff and Lamon (1994) found that
contractions of the L hand led to the expression of negative
emotions such as sadness, whereas contraction of the R hand led to
the expression of positive emotions such as a feeling of well-being.
Because L-pawed animals generally seem to show behavior
patterns consistent with the known functions of the right hemi-
sphere and R-pawed animals show behavior patterns consistent
with the known functions of the left hemisphere, our initial hy-
pothesis was that L-pawed dogs might be more fearful of loud
noises than R-pawed dogs.
Measurement of physiological parameters has also identified
asymmetrical (right hemisphere) control of the hypothalamic-
pituitary-adrenal axis in humans (Henry, 1997; Spivak, Segal,
Mester, & Weizman, 1998), rhesus macaques (Buss et al., 2003),
and rodents (Betancur, Neveu, Vitiello, & Le Moal, 1991; Sullivan
& Gratton, 2002). The sympathetic-adrenomedullary axis is also
considered to be under the dominant control of the right cerebral
hemisphere (Adamec & Morgan, 1994; Wittling, 1997). In gen-
eral, therefore, the right hemisphere is associated with the expres-
sion of intense, often negative, emotions and the control of the
hormonal states that accompany these emotional states (Davidson,
Marshall, Tomarken, & Henriques, 2000).
However, because recent research has described the advantages
of being strongly lateralized compared to weakly lateralized (Rog-
ers, Zucca, & Vallortigara, 2004), we also considered the possi-
N. J. Branson and L. J. Rogers, Centre for Neuroscience and Animal
Behavior, University of New England, Armidale, New South Wales, Australia.
This research forms part of N. J. Branson’s research toward a doctoral
thesis. He gratefully acknowledges support from the University of New
England and also thanks the Kong Company for generously providing Kongs.
We are grateful to S. Cairns for assistance with statistical procedures.
Correspondence concerning this article should be addressed to N. J.
Branson, Centre for Neuroscience and Animal Behavior, School of
Biological, Biomedical and Molecular Sciences, University of New
England, Armidale, New South Wales, Australia, 2351. E-mail:
Journal of Comparative Psychology Copyright 2006 by the American Psychological Association
2006, Vol. 120, No. 3, 176 –183 0735-7036/06/$12.00 DOI: 10.1037/0735-7036.120.3.176
bility that the canine behavioral disorder of noise phobia might be
associated with weak or absent paw preference. The main finding
on which we based this idea was that nonlateralized chicks are
unable to perform two tasks simultaneously (discriminating peb-
bles from grain and being vigilant for a predator), whereas later-
alized chicks can perform both tasks well (Rogers et al., 2004).
The nonlateralized chicks also made more distress calls in re-
sponse to seeing the predator (Dharmaretnam & Rogers, 2005),
indicative of elevated fear responsiveness.
Noise phobia involves the expression of excessive fear in re-
sponse to a sound stimulus (Shull-Selcer & Stagg, 1991). Dogs
with noise phobia are said to display a level of fear that is relative
to the intensity of a sound (Overall, 2002). For instance, low-grade
fear responses might include pacing, panting, and staying close to
the owner; as the intensity of the stimulus increases, the dogs may
exhibit more extreme responses and even sustain physical injuries
while attempting to flee by digging or jumping through glass
windows (Voith & Borchelt, 1985). The phobic response may also
involve freezing for long periods of time (Murphree, Dykman, &
Peters, 1967; Voith & Borchelt, 1985). In fact, there is consider-
able variation in the responses of noise-phobic dogs to particular
sounds (Beerda, Schilder, van Hooff, & de Vries, 1997). Although
no specific data are available on the prevalence of noise phobia in
the domestic dog population, it is not uncommon and it is generally
accepted that all dog breeds and both sexes are susceptible to noise
phobias (Overall, Dunham, & Frank, 2001; Voith & Borchelt,
Of relevance to our study of lateralization and noise phobia,
Watson, Clark, and Tellegen (1988) reported that less lateralized
humans are more prone to experience maladaptive levels of anx-
iety than are more strongly lateralized individuals. There are also
reports of a higher proportion of less lateralized individuals pre-
senting with psychosis (Chapman & Chapman, 1987), schizophre-
nia (Crow, 1997), alexithymia, and post traumatic stress disorder
(PTSD; Parker, Keightley, Smith, & Taylor, 1999; Spivak et al.,
1998; Zeitlin, Lane, O’Leary, & Schrift, 1989). Given that alexi-
thymia and PTSD are associated with the expression of extremely
intense emotions, they may provide a comparison with noise
phobia in the dog, as suggested by Overall (2000) and Thompson
and Shuster (1998). Because human patients with alexithymia and
PTSD have been found to show a higher incidence of ambilater-
ality and that this has been linked to impaired interhemispheric
communication (Parker et al., 1999), we thought that investigating
the relationship between noise phobia and brain lateralization in
dogs might extend our understanding of the relationship between
brain activity and emotional behavior.
We categorized dogs as L-pawed, R-pawed, or ambipreferent
(A) using a new test that avoided the lengthy procedures used in
previous studies (Quaranta, Siniscalchi, Frate, & Vallortigara,
2004; Tan, 1987; Wells, 2003). In addition, we assessed their
reactions to thunderstorms and fireworks.
Access to 48 healthy adult dogs (Canis familiaris; 24 male and 24
female) was obtained through veterinary practices in Geelong, Victoria,
Australia. All of the dogs had been neutered surgically. Their ages ranged
from 2 to 15 years (mean SEM 6.7 1.69 years), and the group was
a mixture of pure and crossbred dogs of small, medium, and large body
Paw Preference Testing
Each dog was visited at its owner’s home by the observer (N. J. Branson)
and presented with a Large Classic Kong, which is a hollow, 10-cm long,
conical shaped, firm rubber tube with a 10-mm hole at one end and a
25-mm hole at the other end (Kong Company, Golden, CO). Although
none of the dogs were specifically food deprived, most had not eaten for
12–18 hr before being presented with the Kong. To encourage paw use, we
filled the Kong with chicken and rice sausage meat and presented it to each
dog on a flat surface. The dog’s use of the L or R forepaw or both forepaws
together (B) to hold the Kong while eating its contents was recorded until
a total of 100 L plus R scores had been collected from each dog (i.e.,
irrespective of the number of B scores). A single score of L or R paw use
was recorded irrespective of how long a paw remained on the Kong. The
intervening events separating scores of paw use included moving the
paw(s) from the Kong and then replacing it on the Kong shortly after it had
been moved from the Kong, and performing another unrelated behavior
between holding the Kong with the paws. If the dog pushed the Kong with
the nose so that it came to rest against a foreleg or if it rolled against a
foreleg, no score of paw use was recorded. To prevent any form of social
reward (verbal or tactile) from affecting the dog’s performance on the task,
the observer ensured no interaction took place during testing.
The average time to collect these data was 30 min per dog. For 36 (75%)
of the dogs, this was achieved in one scoring session. The remaining 12
dogs required a second 30-min scoring session applied a week later. Repeat
measures of 100 paw scores collected 6 months after the first scores were
also recorded for 26 of the dogs in this study, selected randomly from the
total group of 48. The second test was performed to check whether dogs
expressed the same paw preference on both tests.
A third test involving measurement of paw preference on a natural task
was applied to another, randomly selected, subsample of 15 dogs. Scores
were collected while the dog held down a bone while chewing it. This test
was conducted 9 months after the first Kong test. Each dog was presented
with a raw beef bone approximately 30 cm long, and the unimanual (L and
R) and bimanual (B) forepaw use to hold the bone was recorded until a
total of 100 L plus R paw uses had been collected for each dog. As for the
Kong task, L or R paw use was recorded when a paw was placed or
replaced on the bone to hold it in position to allow the dog to chew it.
Questionnaire Rating Responses to Fireworks and
The dog owners were asked to complete a questionnaire to grade their
opinion of each dog’s responses to thunderstorms and fireworks. Owners
were required to respond with either a “yes” or a “no” to questions asking
whether their dog displayed any of the following listed behavioral types:
shaking of the body, dilated pupils, panting, salivating, urinating, defecat-
ing, ears back, corners of the mouth retracted down and back, tail between
the legs, running away, hiding, seeking attention, or barking. Each yes
response was allocated a score of 1, and the total for each dog was used to
generate a response index, henceforth referred to as the questionnaire noise
score. The highest possible score was 13, and the lowest score was 0. All
48 dogs were assessed using this procedure.
Response to Playback of Fireworks and Thunderstorms
An audio recording of the sounds of thunderstorm and fireworks (Sound
Design Studios, 2000) was played to a subsample of 31 dogs (also selected
at random from the larger group) in their home environment. These
experiments were also performed 9 months after the first Kong test. To
balance the procedure for any possible order effect, we played the thun-
derstorm first to 16 of the dogs and played the fireworks first to 15 of the
dogs. Each sound was played for 2 min at a volume of 80–100 db
(measured with a Precision Sound Level Meter, Type 2206, Bru¨el & Kjær,
Nærum, Denmark at 1 m from the speakers in a soundproof room). A
5-min interval was allowed between each playback. The observer (unaware
of the questionnaire noise score at the time) recorded the dog’s response to
the playback using a scoring sheet ranking the responses as listed above for
the questionnaire.
The same subsample of 31 dogs was tested again 2 months after this
experiment in the same manner but this time including a playback of white
noise as well as the fireworks and thunderstorm, the order of presentation
being random. All three sounds were played at an intensity of 80 dbA for
2 min, and there was a 5-min interval between each presentation. This
second test was given to see whether the pattern of responses might remain
the same even though habituation might occur and whether the dog’s noise
reactivity would generalize to white noise.
Experiments were conducted in accordance with the Australian Code of
Practice for the Care and Use of Animals for Scientific Purposes (National
Health and Medical Research Council, 1997) and were approved by the
University of New England Animal Ethics Committee.
Statistical Analysis
The first 100 L or R paw scores were used to calculate a binomial z score
for each dog to determine whether the paw preference differed significantly
from chance. The formula used to calculate this was z (R 0.5N)/
(0.25N), where R signifies the number of R paw uses and N signifies the
sum of L plus R paw uses. Dogs with a positive z score value 1.96 were
R-pawed, those with a negative z score value 1.96 were L-pawed, and the
remainder were ambilateral, A (showing no paw preference). The use of
both paws together was also determined as a percentage of the total number
of unimanual (L plus R) plus bimanual (B) scores. The values for %B were
arcsine transformed prior to statistical analysis to meet parametric
A handedness index (HI) was also calculated for each dog (L R/L
R); hence a score of 1.0 represents exclusive use of the L paw and 1.0
exclusive use of the R paw. The absolute value of HI is the strength of paw
preference with the highest possible value of 1 indicating the exclusive use
of either the L or R paw. The lowest possible HI score is 0 indicating equal
use of the L and R paws.
For all statistical tests, Minitab 13.1 (Minitab Inc., State College, PA)
was used, and the results were considered significant if p .05. A
Ryan–Joiner test on the questionnaire noise scores ( p .10) and a
Levene’s test for equal variance ( p .85) found no evidence to suggest
that the data were not normally distributed. Paw preference classifications
(L, A, or R) and sex were compared with regard to the questionnaire noise
score using the generalized linear model (GLM) procedure for analysis of
variance (ANOVA). Post hoc testing was performed using Tukey’s tests.
The group effect size is given by the statistic omega-squared (w
) for all
fixed-effect ANOVAs and by partial eta-squared (
) for the repeated
measures ANOVAs.
Paw Preference
The z-score calculations of data collected on the 48 dogs tested
on the Kong test identified 21 dogs as being significantly L-pawed,
16 as significantly R-pawed, and 11 as ambilateral. There was a
significant negative correlation between %B and the strength of
paw preference determined when only one paw was used at a time,
r(48) ⫽⫺.44, p .002: Dogs with weaker paw preferences
determined from unilateral scores used both paws together (%B)
more often than dogs with stronger paw preferences.
The alternation of L and R paw use was analyzed using the
Wald–Wolfowitz runs test to identify whether or not each dog’s
use of the L and the R paw was in bouts. No significant runs were
found for any dog ( p .05).
Repeat Paw Preference Testing
To ascertain whether the Kong paw preference was a stable
individual characteristic, we applied a repeat Kong test of paw
preference to a subsample of 26 dogs, 6 months after the first test:
Pearson correlation of the HI scores, r(26) 0.9, p .001 (see
Figure 1A). There was also a positive and statistically significant
correlation between the HI determined in the first Kong test and HI
determined for holding the bone, r(15) 0.96, p .001 (see
Figure 1B). Hence, the measure of paw preference was a stable
characteristic of the dogs.
Relationship Between Paw Preference and Questionnaire
Noise Score
The scores obtained by the questionnaire were analyzed by
GLM with the variables paw preference (L, A, R) and sex. There
was a significant main effect of paw preference, F(3, 47) 4.42,
p .02,
0.1, but no significant effect of sex, F(1, 48) 3.08,
p .09,
.04, and no significant interaction between sex and
paw preference, F(2, 48) 0.55, p .58,
0.08. Post hoc
Tukey testing found a significantly greater questionnaire noise
score for ambilateral dogs compared with L-pawed dogs, t(30)
2.85, p .02, and R-pawed dogs, t(25) ⫽⫺2.53, p .04 (see
Figure 2). There was no significant difference between the scores
for L- and R-pawed dogs, t(35) 0.19, p .98.
The scores for the strength of paw preference were correlated
with the questionnaire noise score, and a significant negative
relationship was found, r(48) ⫽⫺.34, p .02. The correlation
between %B and questionnaire noise score was not significant,
r(48) .11, p .52.
Relationship Between Questionnaire Scores and Measured
Reactivity to Playback of Sounds
The mean reactivity scores determined in the first test in which
sounds (thunderstorm and fireworks) were played correlated pos-
itively with the owner-derived questionnaire noise score, r(31)
0.8, p .001 (see Figure 1C). A GLM analysis with the variables
paw preference (L, R, ambilateral) and treatment (thunderstorm vs.
fireworks sounds, as repeated measures) revealed that paw pref-
erence had a significant main effect on the reactivity to the sounds
of thunderstorm and fireworks, F(2, 30) 9.58, p .001,
0.41 (see Figure 3). Treatment (thunderstorm vs. fireworks)
had no significant main effect, F(1, 30) 0.01, p .75,
0.004, and there was no significant treatment by paw pref-
erence interaction, F(2, 28) 1.0, p .38,
0.07. An a
posteriori Tukey’s test showed that there was no difference be-
tween L- and R-pawed dogs, and that both L- and R-pawed dogs
differed from ambilateral dogs ( p .05).
A significant main effect of paw preference was also identified
in the second playback test, F(2, 29) 6.26, p .006,
(see Figure 3), using the mean score for reactivity to thunderstorm
and fireworks. An a posteriori Tukey’s test identified a significant
difference between the scores for the L-pawed and ambilateral
dogs ( p .001) and between scores of the R-pawed and ambi-
lateral dogs ( p .001) but no significant difference between L-
and R-pawed dogs ( p .59).
The reactivity scores obtained in the second playback test were
also analyzed separately for each sound stimulus using GLM.
There was no significant effect of treatment (thunder, fireworks, or
white noise), F(2, 29) 2.04, p .14,
0.07, and no signif-
icant interaction between treatment and paw preference, F(4,
86) 0.61, p .66,
0.04 (see Figure 4). This suggests that
the dogs responded as much to the white noise as to the sound of
fireworks and the sounds of a thunderstorm. However, this result
may be misleading because the majority of dogs did not respond to
white noise. Those that did respond (6 out of 31) responded quite
strongly: note the high variability of responses to white noise
shown in Figure 4.
There was a positive correlation between the responses given to
fireworks and thunderstorms in the first playback test and the
repeat of this test, r(31) .66, p .001. However, the reactivity
of the dogs was lower in the second test than in the first test:
fireworks, t(31) 3.7, p .001; thunderstorms, t(31) 3.95, p
.001 (see Figures 3 and 4). Nevertheless, the same pattern of
responses was recorded for both playback experiments.
The distribution of paw preferences of the dogs tested was 44%
L, 33% R, and 23% ambilateral. Although there were more
L-pawed dogs in the population, there was no obvious population
bias. This result is consistent with two other studies also showing
no population bias for paw preference in domestic dogs (Quaranta
et al., 2004; Wells, 2003). However, one finding that differs
between these latter two studies and our study was that we did not
find an association between sex and paw preference. Wells (2003)
found that lateralized behavior was sex related, with male dogs
being more inclined to use the L paw and female dogs more
inclined to use the R paw. Quaranta et al. (2004) also found that
male dogs were more likely to be L-pawed, and female dogs
showed a nonsignificant trend to use their R paw. Wells (2003) and
Quaranta et al. (2004) included only sexually entire dogs in their
sample, whereas we tested surgically neutered dogs. Although the
effect of surgical neutering on paw preference has not been inves-
tigated, the pattern of these results suggests that sex hormone
status may be influential.
months for 26 dogs. In both cases, the preferences were determined using
the Kong test. There was a strong positive correlation between the two data
sets, indicating that this measure of paw preference is a stable individual
characteristic. B: Scores for handedness index determined on the first Kong
test and when the dog was eating a bone (N 15). Note the strong positive
correlation. C: Mean reactivity scores determined for each dog in the first
test involving playback of the sounds of a thunderstorm and fireworks
plotted against the reactivity score obtained from the questionnaires com-
pleted by the owners. The correlation is positive and significant.
Figure 1. Data for the significant correlations discussed in the text. A:
Scores for direction of paw preference on two occasions separated by 6
We found that the scores of L and R paw use to hold the Kong
were not influenced by runs or bouts and also that the paw
preference measure was a repeatable individual characteristic.
Strong positive and statistically significant correlations were found
between the paw preference scores on the first and second Kong
tests, separated by 6 months, and also between scores on the Kong
tests and in the test using a bone, demonstrating a likely relevance
to naturalistic behavior.
The paw preferences were associated with reactivity to noise.
Ambilateral dogs had higher scores of reactivity to thunderstorms
Figure 2. Data for the mean (and 95% confidence interval) score of noise reactivity determined from the
questionnaires completed by the owners for each paw preference group: L-paw preferring group (L) is
represented as the white bar, the ambilateral group (A) by the black bar, and the right-paw preferring group (R) by
the gray bar. It can be seen that the reactivity to thunderstorms and fireworks is much higher in the ambilateral group.
Figure 3. The mean (and 95% confidence interval) thunderstorm and fireworks audio playback scores (white
noise is not included) plotted for each paw preference group. Note that the same pattern of greater reactivity in
the ambilateral group (A) compared with the left-paw (L) and right-paw (R) preferring groups is seen for both
playback tests. The reactivity was lower to the second playback compared with the first.
and fireworks, as rated by their owners, than either L- or R-pawed
dogs. Consistent with this, a significant negative correlation was
found between the owner’s score for sound reactivity and the
strength of paw preference. The latter result confirms that, inde-
pendent of the criteria used to group dogs as L-pawed, ambilateral,
or R-pawed, dogs with weaker paw preference show greater reac-
tivity to thunderstorms and fireworks. Hence, the results do not
support the first hypothesis that L-pawed dogs might be more
reactive to these sounds but support our second hypothesis of
greater reactivity in dogs with weaker hemispheric lateralization.
The playback experiments were designed to provide an objec-
tive measure of the dogs’ reactions to acoustic stimuli. Although
differences exist between a playback of a sound recording and the
actual events of thunderstorms and fireworks, this technique has
been used to categorize the reactivity of dogs to sounds (Crowell-
Davis, Seibert, Sung, Parthasarathy, & Curtis, 2003) and to desen-
sitize and countercondition dogs with noise phobia (Overall,
2002). We found a positive and statistically significant correlation
between the owner-rated reactivity scores and the measured reac-
tivity in the playback experiments. Our results for playback
showed a significant association between paw preference and the
scores for reactivity to the sounds of thunderstorms and fireworks.
Ambilateral dogs reacted more strongly to hearing these sounds
than did L- or R-pawed dogs.
The scores in the second playback experiment were lower than
those for the first playback, showing that a degree of habituation
occurred despite the fact that 60 days separated these two tests. In
the only (published) study available for comparison, Crowell-
Davis et al. (2003) did not record any difference between the dog’s
response to a first and second playback of thunderstorm sounds
and fireworks when the presentations were separated by 120 days.
It may be relevant that our playback experiments were carried out
in the dog’s home environment, whereas the study by Crowell-
Davis et al. took place at a veterinary clinic.
Despite the lower reactivity scores in our second playback test,
the relationship between the reactivity score and paw preference
was the same as determined in the first test. Hence, the consistent
relationship identified between paw preference and reactivity to
sound, measured by owner report and by the two playback tests,
provides substantial evidence of weaker paw preference being
associated with greater sound reactivity.
In the first playback experiment, there was no significant dif-
ference between the dogs’ responses to playback of the sound of
either a thunderstorm or fireworks. In the second experiment the
dogs were tested with the sounds of thunderstorms, fireworks, and
white noise, all at the same intensity. Figure 4 shows a nonsignif-
icant trend for the reactivity to playback of the thunderstorm
sounds to be higher than that for fireworks and also for white
noise. Playback for white noise was its first presentation, com-
pared with repeat playback of the other two sounds, which might
suggest that the dogs were less reactive to it than to thunderstorms
and fireworks. Also, the reactivity to white noise was more vari-
able than to the other two sounds: Most dogs made no response to
the white noise but some dogs, particularly in the ambilateral
group, reacted strongly to it. In fact, white noise played at a sound
intensity similar to that used in our experiment has been found to
stimulate the hypothalamic-pituitary-adrenal axis in dogs (Enge-
land, Miller, & Gann, 1990).
We also scored simultaneous use of both paws (%B) to hold the
Kong. A significant negative correlation was found between %B
and the strength of unimanual paw use (absolute value of HI):
Dogs with a weaker preference were also more likely to use both
paws together. However, there was no significant relationship
between %B and reactivity to noise, likely because of the corre-
Figure 4. The mean (and 95% confidence interval) results of the two playback experiments showing the
differences in reactivity to the different sounds. T refers to thunderstorm, F to fireworks, and W to white noise.
The scores for all dogs are presented, irrespective of the paw preference. Note that reactivity to both the
thunderstorm and fireworks is equivalent in playback tests. The dogs’ reactivity was lower in playback Test 2,
and there was no significant difference in reactivity to any of the sounds. Nevertheless, the larger variability to
white noise should be noted: Most dogs did not react to white noise.
lation between %B and strength of paw preference being mild
rather than strong.
The issues we addressed in this study were whether (a) the
presence or absence of lateralization or (b) a lateralized bias to
prefer the L or R paw is associated with the expression of extreme
reactivity to noise, and our results supported the first hypothesis.
As mentioned in the introduction, other studies have identified
behavioral differences between lateralized and nonlateralized in-
dividuals. Of these studies, one of the most relevant for compar-
ison with our results showed that chicks that are nonlateralized for
processing visual information produce more distress calls in re-
sponse to seeing a simulated predator than do lateralized chicks
(Dharmaretnam & Rogers, 2005). It seems possible that nonlater-
alization of neural functions may be associated with intense emo-
tional responses to a broad range of stimuli. One way of inhibiting
an intense emotional response to a disturbing stimulus is to shift
attention to another, less disturbing stimulus, and, from research on
chicks (Rogers et al., 2004), it seems that a lateralized brain is able
to do this more successfully than a nonlateralized brain.
Here we note that dogs exhibit behavioral disorders that may be
homologous to some psychiatric disorders in humans (Overall,
2000; Thompson & Shuster, 1998) and that, as well as showing an
increased incidence of ambilaterality, PTSD patients show a bidi-
rectional interhemispheric transfer deficit (Parker et al., 1999;
Zeitlin et al., 1989). Noise phobia in dogs might depend on a
similar central nervous system mechanism, although absence of
asymmetry at the level of the amygdala may be as important as at
the cortical level, given the role of the amygdala in emotion (Baas,
Aleman, & Kahn, 2004). The antecedent events to noise phobia
and ambilaterality would be interesting to determine because it is
known, in rodents, that early experience (handling and an enriched
environment) has a long-term impact on lateralized brain function
(Denenberg, 1981) and the direction of paw preference (Tang &
Verstynen, 2002).
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Received July 15, 2005
Revision received October 25, 2005
Accepted February 3, 2006
... A major incentive to determine paw preference is that it may be associated with different levels of emotional predispositions and thus may be useful for selecting working dogs. Branson and Rogers found that laterality was associated with greater noise reactivity (Branson and Rogers, 2006); Wells et al. suggested that laterality was associated with "pessimism" and "negativity" in dogs ; and, Barnard et al. report that they "found evidence of a link between canine personality and behavioral laterality, and this was especially true for those traits relating to stronger emotional reactivity, such as aggressiveness, fearfulness, and sociability." . However, other studies have found no relationship between paw preference and emotional predispositions (Wells et al., 2019;Simon et al., 2022a). ...
... Sinescalchi summarized the case for an association between emotions and laterality, specifically stating that left hemisphere dominant dogs (right-pawed) are more aggressive and less fearful than leftpaw dominant or ambilateral-pawed dogs (Siniscalchi et al., 2019). However, other studies have shown that being right-or left-pawed is less significant than the extent of the laterality (Branson and Rogers, 2006). Dogs with a lower laterality index (ambi-pawed) are more fearful and less likely to graduate from seeing-eye school (Batt et al., 2009). ...
... Dogs with a lower laterality index (ambi-pawed) are more fearful and less likely to graduate from seeing-eye school (Batt et al., 2009). There is some evidence that paw preference is consistent throughout the life of a dog (Branson and Rogers, 2006;Batt et al., 2008;Tomkins et al., 2012;Wells et al., 2018). If this is indeed so, then an early indication of the dog's paw preference could aid in the selection of working dogs. ...
... Over the last decades, a substantial increase in research on functional cerebral asymmetries in animals has taken place [1][2][3] . Domestic dogs (Canis lupus familiaris) are one of the key species which have been studied to elucidate the evolutionary and biological mechanisms of lateralization and, thus, to better understand the functional significance of lateralization for vertebrate animals in general 4,5 . ...
... For example, some authors report a relationship between left-pawedness and greater reactivity to stress response, as well as a more pessimistic character in dogs 1,17 . Several different studies showed that measuring the strength of lateralization may also be relevant to understanding the behavioral and emotional characteristics of dogs 2,11,14,18 . For instance, dogs with stronger paw preferences exposed to novel stimuli and unfamiliar environments displayed more confident and relaxed behavior 18 . ...
... For instance, dogs with stronger paw preferences exposed to novel stimuli and unfamiliar environments displayed more confident and relaxed behavior 18 . Weaker lateralization has been identified as a contributing factor to susceptibility to stress, anxiety, fear, and phobia in dogs 2,19 . ...
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Dogs are one of the key animal species in investigating the biological mechanisms of behavioral laterality. Cerebral asymmetries are assumed to be influenced by stress, but this subject has not yet been studied in dogs. This study aims to investigate the effect of stress on laterality in dogs by using two different motor laterality tests: the Kong™ Test and a Food-Reaching Test (FRT). Motor laterality of chronically stressed (n = 28) and emotionally/physically healthy dogs (n = 32) were determined in two different environments, i.e., a home environment and a stressful open field test (OFT) environment. Physiological parameters including salivary cortisol, respiratory rate, and heart rate were measured for each dog, under both conditions. Cortisol results showed that acute stress induction by OFT was successful. A shift towards ambilaterality was detected in dogs after acute stress. Results also showed a significantly lower absolute laterality index in the chronically stressed dogs. Moreover, the direction of the first paw used in FRT was a good predictor of the general paw preference of an animal. Overall, these results provide evidence that both acute and chronic stress exposure can change behavioral asymmetries in dogs.
... Left-, right-or ambilateral preference is characteristic of dogs and cats assessed by scoring the paw used to reach into a stable container to obtain food or, for dogs, to hold steady an unstable container (a Kong) so that they can lick out the food inside [70][71][72]. In a meta-analysis of data on paw preferences in dogs and in cats, Ocklenburg et al. [73] found that most dogs and cats have either a left-or right-paw preference, with no population bias, whereas ambilateral preference is much less common, although the proportion of ambilateral animals varies between samples and the tasks used to score paw preference [74]. ...
... By contrast, other studies of the relationship between paw preference and behaviour have reported that ambilateral dogs differ from both left-and right-pawed dogs in being more reactive to threatening sounds (e.g., the rumbling of a thunderstorm) [71]. Ambilateral dogs are also less aggressive to strangers [78] and show higher levels of fear, playfulness and sociability [79]. ...
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Hand preference in non-human primates has been studied extensively with the aim of understanding the evolution of hemispheric asymmetry and hand preferences in humans. However, the focus has been on hand preferences expressed in adulthood, with a surprising lack of studies on hand preferences in infants and changes that occur during the development of other, potentially associated, asymmetries in the brain and behaviour. This paper reports on the development of hand preference for grasping food and taking it to the mouth in common marmosets. It considers the development of other types of behaviour, such as head cocking and anogenital licking, that parallel and might influence the development of hand preferences during the first months of life. It then discusses behavioural differences between left- and right-handed adult marmosets, including response to novel stimuli, social behaviour and cognitive bias. The need to study the development of hand preferences together with the development of these other expressions of cognitive function is highlighted. The question to be addressed by empirical studies is whether hand preference is a downstream manifestation of the development of hemispheric differences in sensory processing and cognition, or whether it is instrumental in the development of functional differences between the hemispheres. Comparison is made to paw preference and associated behaviour in non-primate species.
... Hand preference in non-human primates has been found to be task specific (McGrew and Marchant 1999), with complexity and animacy of an object affecting hand preference: A right-hand bias has been found only in behaviours directed towards inanimate target objects, but not towards animate targets (Fagot and Vauclair 1991;Forrester et al. 2011Forrester et al. , 2012. Furthermore, the strength of laterality has been shown to be an indicator of emotional functioning in chicks and dogs with weak lateralization being more reactive in response to a stressor (Dharmaretnam and Rogers 2005;Branson and Rogers 2006). It was also found that left handedness is linked to showing stronger fear responses in marmosets (Braccini and Caine 2009). ...
... Handedness, or pawedness, is often assessed by observing the use of an animal's dominant limb when participating in tasks (Batt et al. 2007). Paw preference in dogs has previously been studied through several tasks such as removal of tape on their nose (Quaranta et al. 2004), taking a first step Tomkins et al. 2012; Barnard et al. 2017;Wells et al. 2018), retrieving food from a can (Wells 2003), lifting a paw (Wells 2003;Wells et al. 2018) or stabilising a stuffed Kong™ Branson and Rogers 2006;Schneider et al. 2013;Siniscalchi et al. 2016). From a review of paw preference tests (Wells 2020), the two with the highest test-retest reliability and the easiest to implement for owners, without compromising welfare, are the reaching test (Wells 2003) and lifting a paw test (Wells 2003;Wells et al. 2018). ...
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Handedness has proven to be the most effective and least intrusive measure of laterality in many species. Several studies have investigated paw preference in dogs ( Canis familiaris ) without considering the potential impact that owner’s handedness may have on it, despite dogs being a domesticated species. The aim of this study was to investigate whether owner handedness influences paw preference in their dogs. Sixty-two dogs had their paw preference tested using a Paw Task and a Reach Task in their home over 10 days, recorded by their owners. Interestingly, it was found that left-handed owners were more likely to own a dog with a left paw bias, and right-handed owners were more likely to own a dog with a right paw bias. In the Paw Task, the hand presented to a dog did not significantly predict which paw the dog lifted in response. Furthermore, it was found that females displayed a right paw bias at all age groups. However, males had a left paw bias in puppyhood and right paw bias in older age groups. We conclude that owner handedness influences paw preference in dogs, and it should be considered when suitably pairing dogs to potential owners, especially in assistance work.
... Strength of limb preference can be used as a proxy for the degree of bias to use one hemisphere in preference to using both hemispheres. For example, dogs without a significant paw preference (with an ambilateral preference), measured on the Kong test (see above), react more strongly to the sounds of a thunderstorm than dogs do with left-or right-paw preferences (56). Ambilateral dogs are also found to be both more playful and more aggressive than dogs with significant paw preferences (57). ...
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The specialized functions of each hemisphere of the vertebrate brain are summarized together with the current evidence of lateralized behavior in farm and companion animals, as shown by the eye or ear used to attend and respond to stimuli. Forelimb preference is another manifestation of hemispheric lateralization, as shown by differences in behavior between left- and right-handed primates, left- and right-pawed dogs and cats, and left- and right-limb-preferring horses. Left-limb preference reflects right hemisphere use and is associated with negative cognitive bias. Positive cognitive bias is associated with right-limb and left-hemisphere preferences. The strength of lateralization is also associated with behavior. Animals with weak lateralization of the brain are unable to attend to more than one task at a time, and they are more easily stressed than animals with strong lateralization. This difference is also found in domesticated species with strong vs. weak limb preferences. Individuals with left-limb or ambilateral preference have a bias to express functions of the right hemisphere, heightened fear and aggression, and greater susceptibility to stress. Recognition of lateralized behavior can lead to improved welfare by detecting those animals most likely to suffer fear and distress and by indicating housing conditions and handling procedures that cause stress.
... It seems possible that scarce lateralization of brain functions may be associated with intense emotional responses to a broad spectrum of stimuli. One possible way of inhibiting an intense emotional response to stressful stimulus (i.e. the presence of the unknown human being in our study) would be that of shifting attention to another, less disturbing stimulus, and, from research on vision in chicks 30 and motor lateralization in dogs 31 , it seems that lateralized neural pathways are able to do this more successfully than non-lateralized ones. The antisocial-disruptive behavior of this group could also be partially explained in the light of both "valence" and "approach-withdrawal" models. ...
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There is now scientific evidence that, in dogs, distinctive facial actions are produced in response to different emotionally-arousing stimuli suggesting a relationship between lateralized facial expressions and emotional states. Although in humans, relationships between facial asymmetry and both emotional and physiological distress have been reported, there are currently no data on the laterality of dogs’ facial expressions in response to social stimuli with respect to canine behavioral disorders. The aim of the present work was to investigate the facial asymmetries of dogs with fear and aggressive behavior towards humans during two different emotional situations: (1) while the dogs were alone in the presence of their owners and (2) during the approach of an unfamiliar human being. Overall, our results demonstrated high levels of asymmetries in facial expressions of dogs displaying fear and aggressive behaviors towards humans indicating that measuring facial asymmetries in dogs could prove to be a useful non-invasive tool for investigating physiology-based behavioral disorders.
... Sex effects were detected in dogs' lateralized paw use when retrieving food from a container (Laverack et al., 2021;Wells, 2003), when holding a Kong™ , or when removing an adhesive tape from their head (Quaranta et al., 2004(Quaranta et al., , 2006. However, other studies did not find any sex effects in food retrieving or tape removal tasks Branson and Rogers, 2006;Charlton and Frasnelli, 2022;Wells et al., 2017Wells et al., , 2018. While Tomkins and colleagues (2010) documented breed effects in the context of the Kong™ test, other authors could not confirm this effect (Batt et al., 2008;McGreevy et al., 2010). ...
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SIMON, T., K. Guo, E. Frasnelli, A. Wilkinson and D.S. Mills. Testing of behavioural asymmetries as markers for brain lateralization of emotional states in pet dogs: a critical review. NEUROSCI BIOBEHAV REV XX(X) XXX-XXX, XXXX. Domestic dogs (Canis familiaris) hold a unique position in human society, particularly in their role as social companions; as such, it is important to understand their emotional lives. There has been growing interest in studying behavioural biases in dogs as indirect markers (reflecting lateralized brain activity) of their emotional states. In this paper, we not only review the previous literature on emotion-related behavioural lateralization in dogs, but also propose and apply the concept of evidential weight to previous research. This allows us to examine different hypotheses about emotion-related brain asymmetries (i.e., Right-Hemisphere-, Valence-, Approach-Withdrawal-Hypothesis) on the basis of a “likelihood-ist” concept of evidence. We argue that previous studies have not been able to discriminate well between competing hypotheses and tended to focus on confirmation bias than critically assess different hypotheses; as such there is a strong case for more systematic investigation to pull these theories apart. We present the areas for future research and explain their importance for understanding the emotional lives of dogs.
Domestic dogs differ enormously in both their morphology and behavior. Numerous factors can influence the development and expression of canine behavior and, more generally, determine the success of the pet–owner relationship. This chapter considers the role of nature and nurture in shaping canine behavior. The influence of factors intrinsic to the animal is outlined, focusing on research that has explored the role of breed, sex, and cerebral lateralization in guiding canine behavior and cognitive functioning. The chapter goes on to consider the role of more extrinsic factors that can influence the development of dog behavior, discussing the contribution of early experience, source of acquisition, training techniques, and owner-related traits including personality and attachment style. The article points to the enormous amount of individual variation that exists between dogs and the myriad of factors that can work together to shape the behavior and functioning of the animal we see before us.
Motor lateralization is commonly observed through preferential paw use in dogs and cats. Previous studies have uncovered sex-related differences in paw preference, hypothesizing that these differences may be related to sex hormones. The current study aimed to compare neutered and entire individuals to further investigate whether paw preference is influenced by sex hormones. Dog and cat owners were required to fill in a questionnaire with demographic information such as sex and neuter status of their pets. They then carried out two simple paw preference tasks within their homes: a "reaching for food" task and a "reaching for a toy" task. This study revealed an overall preference among the 272 dogs and 137 cats tested to use their right paw in both tasks. In cats, the degree of paw preference (i.e., regardless of the direction) was significantly influenced by an interaction between neuter status and life stage. Also in dogs, both life stage and an interaction between neuter status and life stage tended to influence the degree of paw preference. Post-hoc power analysis revealed a lack of statistical power, suggesting that future studies using a larger sample size are needed to further investigate potential effects of neuter status on paw preference.
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In recent studies of the structure of affect, positive and negative affect have consistently emerged as two dominant and relatively independent dimensions. A number of mood scales have been created to measure these factors; however, many existing measures are inadequate, showing low reliability or poor convergent or discriminant validity. To fill the need for reliable and valid Positive Affect and Negative Affect scales that are also brief and easy to administer, we developed two 10-item mood scales that comprise the Positive and Negative Affect Schedule (PANAS). The scales are shown to be highly internally consistent, largely uncorrelated, and stable at appropriate levels over a 2-month time period. Normative data and factorial and external evidence of convergent and discriminant validity for the scales are also presented. (PsycINFO Database Record (c) 2010 APA, all rights reserved)
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Common marmosets ( Callithrix jacchus) with a right-hand preference displayed shorter latencies to enter a novel room containing novel structures and objects, touched more objects, and performed more touches and more parallax movements than marmosets with a left-hand preference. These results are consistent with specialization of the right hemisphere (left hand) for fear and negative emotional states and specialization of the left hemisphere (right hand) for approach and positive emotional states. There were no effects of age or sex on any of these behaviors. This relationship between exploration and hand preference did not occur when the marmosets were tested in the home cage with a novel problem, although significant effects of both age and dominance were found in solving the problem. (PsycINFO Database Record (c) 2012 APA, all rights reserved)
Objective: Previous research has demonstrated an association between alexithymia and a deficit in interhemispheric communication in Vietnam combat veterans with posttraumatic stress disorder. The purpose of this study was to evaluate this association in a nonclinical sample. Methods: The efficiency of interhemispheric transfer was assessed in 14 alexithymic and 15 nonalexithymic right-handed, male, undergraduate university students using a tactile finger localization task. Results: The nonalexithymic subjects were significantly more efficient at transferring information between the cerebral hemispheres than the alexithymic subjects. Conclusions: This finding provides further evidence of an interhemispheric transfer deficit in alexithymia and suggests that an alexithymic cognitive style reflects poor integration of the information processing of the two cerebral hemispheres.
Twenty-three university students enrolled in an introductory Army Reserve Officers Training Corps (ROTC) course described thirteen aspects of themselves by choosing terms from an idiographically generated list of traits. Among the self-images described were the reflected self-image in the ROTC (the way others see me now in the ROTC), actual career self, and the desired career identity. Each participant's data was individually analyzed using HICLAS. Discrepancies between the reflected self-image and (a) the actual career self and (b) desired career identity were scored from each participant's HICLAS output. Both discrepancy scores predicted intention to continue in the ROTC, assessed six months after the self-descriptive data were collected. The reflected self-image/actual career self discrepancy was the better predictor of intention to continue, and was the only discrepancy score to predict whether or not students actually remained in ROTC in the following academic year. The actual self may be the more reliable reference point to which the reflected self image is compared, and the discrepancy between these images of self may inform people's judgment of whether they fit in with a group.
Among right-handers, the magnitude of differences in proficiency between the left and right hands varies considerably. Yet significance of the extent of right-handedness is still a controversial issue. To examine whether individual differences in asymmetry of hand skill can partly be attributed to individual differences in asymmetrical hemispheric activation, handedness and electroencephalographic (EEG) laterality were correlated in two large samples (ns=60 and 128). Analysis indicated thar part of the variability in right-handedness may arise from activation asymmetries in the cortex, but whether this relation becomes apparent depends on the cortical area examined and on the experimental condition under which the EEG measures are taken.
Poor housing conditions, harsh training sessions and uncontrollable or unpredictable social environments are examples of the situations that may lead to reduced welfare status in dogs. Individuals that suffer from poor welfare presumably experience stress and may consequently exhibit stress responses. In order to evaluate stress responses as potential indicators of poor welfare in dogs, we review studies dealing with dogs subjected to stressors. The reported stress responses are categorized as being behavioural, physiological or immunological, and demonstrate the various ways stress is manifested in the dog.
A review of research with chicks, songbirds, rodents, and nonhuman primates indicates that the brain is lateralized for a number of behavioral functions. These findings can be understood in terms of three hypothetical brain processes derived from a brain model based on general systems theory: hemispheric activation, interhemispheric inhibition, and interhemispheric coupling. Left-hemisphere activation occurs in songbirds and nonhuman primates in response to salient auditory or visual input, or when a communicative output is required. The right hemisphere is activated in rats when spatial performance is required, and in chicks when they are placed in an emotion-provoking situation. In rats and chicks interhemispheric activation and inhibition occur when there is an affective component in the environment (novelty, aversive conditioning) or when an emotional response is emitted (copulation, attack, killing). An interhemispheric coupling (correlation) found in rats and rabbits implies that the hemispheres are two major components in a control system with a negative feedback loop. Early-experience variables in rats can induce laterality in a symmetric brain or facilitate its development in an already biased brain. It is predicted that functional lateralization, when present, will be similar across species: the left hemisphere will tend to be involved in communicative functions while the right hemisphere will respond to spatial and affective information; both hemispheres will often interact via activation-inhibition mechanisms when affective or emotional processes are involved. Homologous brain areas and their connecting callosal fibers must be intact at birth and must remain intact throughout development for lateralization to reach its maximum level. Injury to any portion of this unit will result in hemispheric redundancy rather than specialization. One major function of early experience is to provide stimulation during development, which acts to enhance the growth and development of the corpus callosum, thereby giving rise to a more specialized brain.