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Paw Preferences in Dogs

Üner Tan a
a
Atatürk University, Medical Faculty, Institute of Physiology, Erzurum, Turkey

Online Publication Date: 01 February 1987

To cite this Article Tan, Üner(1987)'Paw Preferences in Dogs',International Journal of Neuroscience,32:3,825 — 829
To link to this Article: DOI: 10.3109/00207458709043336
URL: http://dx.doi.org/10.3109/00207458709043336

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 Infern. J . Neuroscience. 1987. Vol. 32, pp. 825-829 0 1987 Gordon and Breach, Science Publishers, Inc.
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PAW PREFERENCES IN DOGS

UNER TAN
Atu turk University, MedicoI Fu cu 18, lnstitu t e of Physiology, Erz u rid mi, Ti1rk ey

( Keceiveti Mriy 22, 1986)

T h e distrihution o f paw preferences were studied in 2X dogs. T h e paw preference was assessed by
counting the right and left paw movements performed to remove an adhesive plaster from the eyes. The
significance o f the right minus left paw reaches in percentages was evaluated statistically in each animal.
There were threc distinct groups in rcspcct to paw preferences in dogs: right-preferent (57. I”/”), left-
preferent ( 17.9‘%,),and ambidextrous (25.0%). Statistical analysis showed that the observed frequencies
for each group were not merely chance variations which would be expected in a random sample. It was
concluded that the population bias can he expressed in a distrihution skewed toward a right-hand bias as
seen in man.
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hrind preJerence, po I+’prejkrence, dog

Keywords: L(iterrilbation, (isymmel~y,

In 1862, Buchanan stated that “the use of the right hand in preference to the left
must be regarded as a general characteristic of the family of man.” Since then, this
assertion does not seem to be changed essentially, but the origin of human
handedness remained obscure. The attempt was made to approach the problem by
animal studies, which, however, did not give any satisfactory results, since the paw
preferences among animals were found to be equally distributed in the population.
In chimpanzees (Finch, 1941), orang-utans (Annett, 1985), rhesus monkeys
(Warren et al., 1967),cats (Cole, 1955),rat (Tsai & Maurer, 1930; Peterson, 1934),
and mice (Collins, 1969), approximately 50% of animals showed ambidextrality,
and the remainder were about equally divided between right- and left-preferents.
Thus, it seems that there is no species bias to the right side in any mammal other
than man, and the handedness in nonhuman mammals seems to be a stochastic
process. However, as stated by Annett ( 1989, the consistency of hand preferences
varies between tasks and the animal’s experience of a task. On the other hand,
animal studies would be essential to understand the origin of human handedness.
For these reasons, it still appears to be of considerable importance to study
asymmetrical animal behavior such as handedness. Accordingly, paw preferences
were studied in dogs, which remained unattended in the field of motor
lateralization. The results suggest that the distribution of paw preferences in dogs
show some similarities to the distribution of hand preferences in man.

METHODS

The animals were 28 adult mongrel dogs collected from villages 19 of which were
female, and 9 male. The animals had no other behavioral testing before or during
hand testing. To assess paw preference, the eyes of an animal were closed by an
adhesive plaster. The dog was then allowed to try removing it from the eyes by
using either the right or left paw. The frequency of paw use was evaluated by
825
 826 UNER TAN

pressing a counter at each paw movement. Two counters were used, one for the
right and one for the left paw. This procedure was continued until the sum of the
right and left paw reaches was 100. Only intentional, goal-directed paw movements
were taken into account. In each animal, the significance of the difference between
the frequencies of the right and left paw movements was assessed by the binorninal
test for large samples. According to the significance of the difference between the
two frequencies, the animals were designated as right-preferent, left-preferent, or
ambidextrous. The significance of the difference in the group frequencies (number
of animals in each handedness group) was tested by the Kolmogrov-Smirnov one-
sample test.

RESULTS

Figure 1 shows the distribution of the right minus left paw movements carried out
to reach a target in 28 dogs. The sum of the right and left paw reaches was taken as
100 in each animal. The difference between the right and left paw reaches was
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significantly larger than zero in 16 animals, and smaller than zero in 5 animals.
There was no statistically significant difference between the right and left paw
reaches in 7 animals. Accordingly, the dogs studied for paw preferences constituted
three groups: right-preferent (57.1%), left-preferent ( 17.9%), and ambidextrous
(25.0%). The corresponding three peaks are recognizable in the histogram, which
are designated as L, A, and R, indicating the most commonly occurring right-left
differences within the left-preferent, ambidextrous, and right-preferent groups,
respectively.

( R - L ) Paw use
FIGURE 1 The distribution of paw preferences in dogs ( N = 2 8 ) f o r right minus left p ; ~ wre;ichr it) il
total of 100 right plus left paw movements.

When the left-preferent and ambidextrous animals were considered as a single

group, ix., non-right-preferent group ( N = 12, 42.9%), and the right-preferent
animals as another group ( N = 16, 57.1%), the probability of the former ( p , )and the
probability of the latter group ( p ? )was found to be statistically equal, that is,
p , = 17. = ( z = - 3 7 , p = .28). It could be concluded that the difference between the
 PAW PREFERENCE IN DOGS 827

two groups arose because of chance. However, considering the magnitude as well
as the direction of the differences between the total right, and left paw use in each
animal, it was found that the sum of the negative scores (right minus left paw use
smaller than zero) was significantly less than the sum of the positive scores (right
minus left paw use larger than zero). The Wilcoxson matched-pairs signed-ranks
test for large samples was used to test this difference. At T= 106, z= - 1.61, which
was significant at p = .05 (alpha = .05). Thus, the number of animals using the right
paw more frequently than the left paw significantly exceeded the number of animals
using the left paw more frequently than the right paw, and the dogs' right paw use
was significantly more frequent than the left paw use in the total sample.
The frequency of the left-preferent animals ( N = 5 , 17.9%) is significantly
different from the sum of the right-preferent and ambidextrous animals ( N = 23,
82.1%) at p=.0007 ( z = -3.21). The frequency of the ambidextrous animals
( N = 7 , 25.0%) was also significantly less than the' frequency of the remainder
( z = - 2.46, p = .007). Thus, the difference between the left-preferent or
ambidextrous dogs and the sum of the others did not arise because of chance. The
binominal test for large samples was used to determine the significance of the
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difference between the frequencies of the above groups because the data could be
considered in two discrete categories.
The Kolmogrov-Smirnov one-sample test was used, to determine whether the
frequencies of the right-preferent, left-preferent, and ambidextrous dogs were
distributed equally. The maximum deviation was found to be .24, which cannot be
interpreted on the basis of chance ( p = . O 5 ) . Thus, one could conclude that the
animals showed significant preference classes. This result was supported by the x'
one-sample test (X' = 7.38 > zZt; .05 = 5.99).

DISCUSSION

It was statistically shown, in the present work, that there are three distinct groups of
dogs in respect to paw preferences: right-preferent ( 57.1%), left-preferent ( 17.9%),
and ambidextrous (25.0%). Of all the animals, 75% exhibited significant paw
preferences. The frequency of the right-preferent animals did not differ statistically
from the frequency of the remainder, indicating a distribution on the basis of
chance. However, the number of animals showing the right minus left paw use
greater than zero (R - L> 0) significantly exceeded the number of animals showing
R - L < 0; the sum of the paw reaches of R - L < 0 was significantly less than that of
R - L< 0. Thus, there was a tendency to bias to the right paw use in the total
sample.
The nonparametric statistics showed that the observed frequencies for each
group of paw preferences were not merely chance variations, which would be
expected in a random sample from the rectangular population where =f?=&.
Moreover, the frequency of the left-preferents or the frequency of the
ambidextrous animals was significantly less than the frequency of the remainder,
These findings do not support the notion that handedness in nonhuman mammals
merely depends on chance, which was inferred from the common finding that there
are as many animals left-preferent as right-preferent, and about 50% of the animals
are mixed handers.
Annett (1985)analyzed hand and paw preferences in nonhuman mammals and
concluded that the distribution of hand preferences in other species than man is
 828 UNER TAN

just as expected from a normal distribution of the right-left differences with a mean
difference of 0, or L=R. As a substantial difference between human and
nonhuman samples, she suggested that the distribution of hand preference in man
also fits a normal distribution but with a mean to the right of zero; i.e., only the
location of the mean is different in the distribution of human and nonhuman hand
preferences. The present work does not support these theoretical considerations on
the distribution of hand preference of nonhuman mammals. In contrast, it shows a
rather similar distribution of paw preferences in dogs with a shift to the right of
zero as in man. Respectively, Bianki et al. (1979)have reported that there is a right
dominance in hybrids of the domestic mouse, and a left dominance in the
regulation of the general motor activity in white mice.
It was argued that the left-handers do not constitute a distinct group in the
population, considering handedness as a continuous variable (Annett, 1985). It was
found, in the present work, that the left-preferent dogs constitute a distinct group,
and the distribution of paw preference in these animals did not support a unimodal
model. McManus (1989, and Tapley and Bryden (1985) suggested a symmetric
bimodal distribution of hand skill in man.
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The incidence of left-preference in dogs (17.1%) is not very much different from
that. found in man, which ranges from 1 to 30% according to Hecean and
Ajuriaguerra’s ( 1964) review. In general, however, the incidence of left-handedness
in human societies is much lower than that in animal populations. Annett (1YH5)
suggested a theoretical distribution of handedness in man as 66% right-preferent,
30% ambidextrous, and 4% left-preferent. Using the peg moving task, I observed
57.7% right-preferent, 37.1% ambidextrous, and 5.2% left-preferent subjects (Tan,
1985). The direction of these proportions is the same as the dogs’ data, but the
proportions are different in man and dog. The reason may be the method used to
assess the handedness. The same is also true for human studies. For instance, when
handedness was determined by performing diverse activities such as writing and
throwing, a J-shaped distribution was obtained with about 90% of the population
showing right-hand preference (Annett, 1972). In contrast, a normal distribution
was obtained by using a peg moving task to compare the relative proficiency of the
two hands (Annett, 1985). Using a paper-and-pencil test, Tapley and Bryden
(1985)found that the distribution of hand performance was clearly bimodal.
The method used to assess the handedness is also important in animal studies.
Kliiver (1933),testing the hand preference in a Java monkey, suggested ‘‘. . . that in
testing for handedness we must keep in mind the possibility that a change in
distance may lead to a change in hand preference.” Sanford et al. (1984)found, in
bushbabies, that the population could be expressed by a distribution skewed
toward a left-hand bias, in the vertical stance, but the hand preference shifted to a
bimodal distribution with equal probabilities in a quadripedal stance. Thus, the
upright position influences the lateralization of the freed hands, and one of the
reasons of less left-handedness in man than dog might be the bipedalism of humans
favoring the hands for a variety of complex actions.

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 PAW PREFERENCE IN DOGS 829

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