Intern. J.

Neuroscience, 113:1675–1689, 2003

Copyright  Taylor & Francis Inc.
ISSN: 0020-7454 / 1543-5245 online
DOI: 10.1080/00207450390249258

POPULATION-LEVEL RIGHT-PAW
PREFERENCE IN RATS ASSESSED
BY A NEW COMPUTERIZED
FOOD-REACHING TEST

MUSTAFA GÜVEN
Department of Biophysics
Medical School, Çukurova University
Adana, Turkey

DERYA DENIZ ELALMIŞ

SEÇIL BINOKAY
ÜNER TAN
Neurophysiology Unit, Department of Physiology
Medical School, Çukurova University
Adana, Turkey

We re-studied the distribution of paw preference in rats using a new

computerized food-reaching test, which recorded the times and time
intervals between the single right- and left-paw entries. Using the tradi-
tional food-reaching test, we found that of 144 rats, 72.7% were right-
handed, 19.7% left-handed, and 7.6% mixed-handed. This population-
level J-shaped right-hand preference did not fit a binomial chance dis-
tribution (25:25:50). Of right-handers, 99.5% first used their right paw
and 0.5% left paw; of left-handers, 98.6% first used their left paw and
1.4% right paw. Of mixed-handers, 59% first used the right paw and
41% left paw for food reaching. The time interval between putting the
rat into the test cage and the first right-paw entry was significantly
shorter than the first left-paw entry in total sample. Males were faster
than females (shorter time intervals between right- or left-paw entries).
The distribution of the time intervals between right- or left-paw entries

Received 30 May 2003.

Address correspondence to Prof. Dr. Üner Tan, Head of the Neurophysiology Unit, Department
of Physiology, Medical School, Cukurova University, Adana, Turkey. E-mail: unertan@cu.edu.tr.

1675
1676 M. Güven et al.

was inverse J-shaped, which exhibited a normal distribution after tak-

ing the logarithms of the time intervals. There was no significant differ-
ence between time intervals for the left-paw entries; time intervals for
the right-paw entries were significantly shorter in males than females,
accentuating the role of the left brain for sex differences in motor con-
trol. The results suggested that humans are not unique in population-
level right-hand preference; our new method would be suitable for new
developments in handedness research.
Keywords food reaching, handedness, laterality, paw preference, paw skill

Is the human being unique in population-level right-handedness?

There is a considerable debate about this subject in the scientific
literature. Until recently, there was a general tendency towards the
uniqueness of man in handedness (see for a review, Elalmis, Ozgunen,
Binokay, Tan, Ozgunen, & Tan, 2003). This consensus was, how-
ever, challenged in 1987 by MacNeilage, Studdert-Kennedy, and
Lindblom; there is now accumulating evidence for population-level
asymmetries in animals, suggesting that this is not exclusively due
to humans (see Bisazza, Rogers, & Vallortigara, 1998). There are
indeed numerous studies with evidence for the presence of popula-
tion-level right-handedness in animals (see for a review, Elalmis et
al., 2003).
In rats, a population-level right-handedness was reported since
1930 (see Elalmis et al., 2003; Glick & Ross, 1981; Pence, 2002;
Tang & Verstynen, 2002; Tsai & Maurer, 1930; Waters & Denenberg,
1994; Whishaw, Pellis, & Gomy, 1992). “There are structural and
functional asymmetries at the population level in fish, amphibians,
and reptiles” (Bisazza, Rogers, & Vallortigara, 1998). Similar to
humans, the hand preference in rats is controlled by the contralat-
eral primary motor cortex, since handedness is reversed after abla-
tion of this region (Castro-Alamancos & Borrell, 1993). Recently,
Elalmis et al. (2003) have reported a population-level right-paw
preference in rats, providing further strong support for the argument
that humans are not unique in hand preference and that a homology
exists between human and rat reaching movement (see Whishaw et
al., 1992). There are, however, some other reports indicating no
population-level right-handedness in rats (Collins, 1985; Otmakhava,
1989; Whishaw et al., 1992). According to Tang and Verstynen
 Paw Preference in Rats 1677

(2002), “these seemingly inconsistent results in the literature can be

explained in terms of the differences among testing methods.”
Elalmis et al. (2003) have recently used a different method for
assessing hand preference in rats. They evaluated the frequency of
paw reaching, the number of the paw entries within 10 min, instead
of simple counting of paw entries without considering the time, and
found that the distribution of paw preference in rats is J-shaped, and
there is a right-sided population bias in handedness as in humans. In
the present work, the time intervals between the consequent paw
reaches were measured using a new computerized method instead
of a global paw frequency (numbers of paw reaches in a limited
time period).
The estrus cycle was determined after testing for hand prefer-
ence, since the estrus cycle was shown to influence the asymmetric
cerebral functions in rats (see for instance, Robinson, Camp, Jacknow,
& Becker, 1982); the inconsistent results concerning the paw pref-
erence of rats may also be accounted for by fluctuations in sex
hormone levels during the estrus cycle. In accord, it was recently
shown that the estrus cycle was a significant factor influencing the
right- and left-paw entry scores in female rats (see Elamis et al.,
2003). Therefore, we have also determined the phase of the estrus
cycle by examining the vaginal smears, which were taken after test-
ing for paw preference, to avoid any effect of a stressor on hand
preference (see Fitzgerald, Glick, & Carlson, 1990).

MATERIALS AND METHODS

Animals

The experiments were performed on male and female Wistar rats

adapted to the animal room for at least two weeks. The animals
were housed in groups of 3–4 and maintained on a 12-h light–dark
cycle. Before testing for paw preference, the rats were deprived of
food for two days and then individually placed in a metal housing
cage for testing. On the test day, the rats were first weighed; it was
required that they 1ose 15% of their initial weight after being food
deprived for two days.
1678 M. Güven et al.

Assessment of Estrus Cycle in Female Rats

Stress is reported to be primarily responsible for the marked right-
sided population bias in male rats during T-maze learning (Fitzgerald,
Glick, & Carlson, 1990). To avoid such a bias in our rats, vaginal
smears were taken from the vulva for the evaluation of the estrus
cycle, immediately after food-reaching test. The vagina was washed
with saline and a small smear was taken into the tip of an Eppendorf
tube. The smear was then put on the surface of a microscope glass,
stained with methylene blue, and examined under microscope. The
vaginal cytology can be used as an endocrine assay, especially for
estrogen. There is a marked increase in blood estrogen levels during
proestrus. During diestrus, the blood estrogen concentration is low
but gradually increases; during proestrus, the blood estrogen con-
centration markedly increases, decreasing during estrus, which is
associated with high progesterone levels. These estrous stages can
be recognized from vaginal smears.

Paw Preference
The paw preference of rats was essentially assessed by the method
used by Tang and Verstynen (2002): there were two opening in the
front of the testing cage, separated by 1 cm. These openings were
small enough to allow access to food rewards by the forepaws, not
by the snout. Rats were classified as right-, left-, and mixed-handers
based on the binomial probability distribution test (see Betaneur,
Neveu, & Moal, 1991): rats were considered right-pawed if the right-
paw entry (RPE) score was equal to or greater than 29, left-pawed if
the left-paw entry (LPE) score was equal to or smaller than 21, and
ambidextrous when the RPE score was between 22 and 28. The paw
preference was followed for successive 6 days; only the results for
the 5th day were analyzed in the present work, since the paw pref-
erence stabilized during this time (see Elalmis et al., 2003). The
animals were weighed every day to control the daily loss of the
weight, which should not exceed 15% of the original body weight.
A computer was used for recording the fast and accurate times of
reaching movements, using computer-software, which was referred
by us as “Activity Logger.” This software was written in C++ pro-
gramming language and was run under Windows98 operating system.
 Paw Preference in Rats 1679

The software included a time counter in 1 ms precision, logging

windows, buttons, and menus. When the experiment was started
after putting the rat into the test cage, the time counter was started
by pressing the “start button.” At each paw entry, the appropriate
buttons for the right- and left-paw entries were pressed on the key-
board of the computer. During this procedure, the time counter con-
tinuously recorded the times between paw entries. In doing this, we
could record the time intervals for the right- and left-paw entries. At
the end of the experiment, the data were saved as a text file, which
included four different columns: RPE (the times for the consequent
RPEs), LPE (the times for the consequent LPEs). The saved data
were then analyzed using a statistical package, SPSS for Windows,
V 10.0.1

Statistical Analysis
SPSS for Windows (V. 10) was used for the statistical analysis of
data. Eta squared, the proportion of the total variability in the de-
pendent variable that is accounted for by variation in the indepen-
dent variable, was calculated for each significant ANOVA result.
After recording the times for each paw entry, the instantaneous fre-
quency was calculated by taking the reversal of the time interval
between paw entries (sec–1). A low value of the instantaneous fre-
quency indicated slow paw reaching, and high value corresponded
to fast paw reaching following a previous paw entry.

RESULTS

Distribution of Global Paw Preference

Table 1 presents the numbers and percentages of the male and fe-
male right-, left-, and mixed-pawed rats. There was no significant
sex difference between the relative numbers of the right-, left-, and
mixed-handers (χ2 = 4.6, df = 2, p > .05).

1. For detailed information concerning the software, Dr. Mustafa Güven can be contacted
(guvenm@cukurova.edu.tr).
1680 M. Güven et al.

TABLE 1. Numbers and percentages of right-, left-, and mixed-handed male and
female rats

Right-handers Left-handers Mixed-handers

Rats N % N % N %

Total 144 72.7 39 19.7 15 7.6

Females 96 72.2 30 22.6 7 5.3
Males 48 73.8 9 13.8 8 12.3

In the total sample, all categories (right-handers, left-handers, and

mixed-handers) were not equally distributed (χ2 = 142.7, df = 2, p <
.001). The binomial test rejected the 0.50 test proportion between
right- (N = 144, 79.0%) and left-handers (N = 39, 21.0%), after
excluding the mixed-handers (p < .001, based on Z approximation).
The frequencies of the right-, left-, and mixed-handed rats did also
not fit to 25:25:50, expected from a binomial distribution (χ2 =
253.9, df = 2, p < .001).
Considering strongly right-handed rats (RPE > 45), strongly left-
handed rats (RPE < 5), and mixed-handers (RPE = 5 to 45), of 197
rats, 70 (35.5%) were strongly right-handed, 9 (4.6%) were strongly
left-handed, and 118 (59.9%) were mixed-handed. There was no
significant sex difference in the numbers of these subgroups (χ2 =
5.1, df = 2, p > .05), although the numbers of the strongly right-
handed (N = 52/133, 39.1%) and strongly left-handed (N = 8/133,
6.0%) females exceeded the number of the strongly right-handed
(N = 18/64, 28.1%) and strongly left-handed (N = 1/64, 1.6%) males;
there were more mixed-handed males (N = 45/64, 70.3%) than fe-
males (N = 73/133, 54.9%).
Figure 1 illustrates the distribution of the RPE scores per 50
trials in the male (B) and female (A) rats. The distribution of paw
preference in rats seemed to be J-shaped, neither U-shaped nor nor-
mal. The preponderance and accumulation of the right- and left-
handed female rats and mixed-handedness in male rats are visible in
Figure 1A and 1B, respectively.
Although the weight of the rats was kept constant for each test-
ing day, any correlation between weight and RPE scores was exam-
ined to be certain about the effects of weight on food reaching. The
 Paw Preference in Rats 1681

FIGURE 1. Distribution of right-paw entry scores (right + left = 50) in the female (A)
and male (B) rats. Abscissa: RPE score; ordinate: percentage of rats.

statistical analysis indicated that there was no significant correlation

between these parameters (r = –.14, N = 196, p > .10).

First Paw Used for Food Reaching

In right-handed rats, of 2032 trials the number of the RPE was 2022
(99.5%) first used to reach the food, there were only 10 (0.5%)
LPE first used for food reaching. The left-handed rats exhibited an
1682 M. Güven et al.

opposite distribution: Of 682 RPEs, only 10 (1.4%) paws were the

right paw first used for food reaching; 672 (98.6%) left paws were
the first to reach the food. Of 137 first-paw reaches, in mixed-
handed rats, 82 right-paw (59%) was the first and 55 (41%) left-
paw was the first to reach the food.In these mixed-handed rats, the
number of the right-paw first reached was significantly greater than
the number of the left-paw first reached (χ2 = 9.9, df = 1, p < .005).

Times for the First RPEs and LPEs

Total Sample
Table 2 presents the mean, standard deviation, median, minimum,
maximum, 25th percentiles, 50th percentiles (median), and 75th per-
centiles for the first time interval (s) between putting the rat into the
test cage and the first RPE and LPE, in the total sample. The mean
time interval for the first RPE was significantly shorter than the
mean time interval for the first LPE (Wilcoxon Signed Rank Test:
z = –5.67, p < .001).

TABLE 2. Mean, standard deviation, minimum, maximum, 25th, 50th (median), and
75th percentiles for the times (s) elapsed between putting the rat into the test cage and
the first RPE or LPE in rats

Samples N Mean Std. Min. Max. 25th 50th 75th

Total
RPE 57 101.9 193.7 1.33 820.3 6.62 18.11 81.89
LPE 48 307.1 307.1 7.59 1322.5 33.91 103.89 283.52
RPE
Males 33 99.3 200.6 1.33 773.3 4.33 14.41 95.59
Females 46 229.7 290.5 1.83 1415.2 39.56 137.99 419.22
LPE
Males 30 161.0 205.2 9.96 773.6 19.86 61.49 259.63
Females 41 338.9 360.7 8.23 1328.5 78.12 200.34 537.12
RHs
RPE 275.4 361.7 1.33 1536.0 27.17 97.76 396.99
LPE 370.0 381.3 7.60 1849.4 100.39 201.07 469.13
LHs
RPE 565.9 392.1 32.0 1703.4 218.03 485.58 896.86
LPE 427.2 379.0 5.3 1476.2 97.88 247.57 754.02
MHs
RPE 597.4 301.7 1.86 1197.7 361.11 629.90 762.86
LPE 642.3 335.1 9.96 1330.9 377.18 648.85 889.63
 Paw Preference in Rats 1683

Sex Differences
The mean time for the first RPE was significantly shorter in males
than females (Mann-Whitney Test: z = –17.7, p < .001). The 25th,
50th, and 75th percentiles for the first RPE in males were also less
than females (see Table 2).
The mean time for the first LPE was significantly shorter in males
than females (z = –12.56, p < .001). The 25th, 50th, and 75th per-
centiles in males were also less than those in females (see Table 2).

Right-, Left-, and Mixed-Handers

As expected, the time for the first RPE was significantly shorter
than the time for the first LPE in right-handed (RPE > 28) rats
(Wilcoxon Signed Rank Test: z = –17.06, p < .001). Accordingly,
the percentile values were also shorter in males than females (see
Table 2).
In left-handed rats, the mean time interval for the first LPE was
significantly shorter than that for the first RPE (Wilcoxon Signed
Ranks Test: z = –8.82, p < .001). The percentile values for the first
LPE were also less than the first RPE (see Table 2).
In mixed-handed rats (RPE, there was no significant difference
between the time intervals for the first RPE and the first LPE (see
Table 2; Wilcoxon Signed Rank Test: z = –0.42, p > .65).

Distribution of Time Intervals Between

Consequent Paw Reaches
Figure 2 illustrates the distribution of time intervals between conse-
quent paw entries for the right (A) and left (B) paws. One-sample
Kolmogorov Smirnov Test indicated that the time intervals between
the consequent right- and left-paw reaches did not fit normal distri-
butions (A: right paw, z = 989, p < .001; B: left paw, z = 7.43, p <
.001). They seemed to be inverse J-shaped distributions. However,
the distributions were normal after taking the logarithms of the time
intervals (C: RPE, z = 1.27, p > .05; D: left paw, z = 1.02, p > .20).
Table 3 presents the means and median values of the time inter-
vals (s) between the consequent paw entries in the total sample,
males, and females. As seen in Table 3, the mean and median values
1684 M. Güven et al.

FIGURE 2. Distribution of time intervals between consequent paw entries for RPE (A)
and LPE (B) in total sample. Abscissa: time interval (s); ordinate: number of RPE in A
and number of LPE in B. RPE (C) and LPE (D) above histograms after logarithmic trans-
formations of the time intervals (C: RPE, D: LPE).

for the time intervals were significantly greater for the left-paw
entries than the right-paw entries (Wilcoxon Signed Rank Test: z =
4.77, p < .001): the times between the single paw reaches were
significantly shorter between the right-paw reaches than the left-
paw reaches in the total sample. After taking the logarithms of the
time intervals, to obtain a normal distribution and use parametric
statistics instead of nonparametric statistics, the mean time interval
between the right-paw entries was found to be significantly longer
than the mean time interval between the left-paw entries, for the
total sample (t = 9.91, df = 3186, p < .00l).
 Paw Preference in Rats 1685

TABLE 3. Mean and median values of time differences between consequent RPEs in
total samples, male and female rats

INTERVAL Log INTERVAL

Number INTERVAL LogINTER
Rats of paws Mean SD median Mean SD median

RPE (TOT) 2387 6.5 7.3 3.85 0.605 0.42 0.586

LPE (TOT) 1065 11.3 14.6 5.62 0.779 0.49 0.752
RPE (M) 1148 5.1 4.8 3.38 0.538 0.38 0.532
RPE (F) 1239 7.7 8.8 4.50 0.666 0.44 0.653
LPE (M) 223 14.3 18.2 6.29 0.824 0.57 0.798
LPE (F) 842 10.5 13.4 5.49 0.768 0.47 0.736
RPE (RH) 2247 6.0 6.6 3.72 0.588 0.41 0.572
RPE (LH) 140 13.1 12.5 8.93 0.870 0.51 0.951
LPE (RH) 373 15.2 19.4 6.63 0.848 0.57 0.822
LPE (LH) 692 9.2 10.6 5.14 0.743 0.44 0.711

The mean time interval between the consequent right-paw reaches

were found to be significantly shorter in males than in females (Mann-
Whitney U Test: z = 6.83, p < .001). The mean log10 RPE-time
interval was also significantly shorter in males than females (F1, 2385
= 56.95, p < .001). With regard to the sex difference in the mean
LPE-time intervals, the mean LPE-time intervals was shorter in fe-
males than males (see Table 3) but the difference was statistically
nonsignificant (Mann-Whitney U Test: z = 1.39, p > .15). The loga-
rithmic scale for the LPE-time intervals did also not reveal any
significant sex difference (F1, 1063 = 2.36, p > .10).
Concerning the handedness for the RPE-time intervals, the right-
handers (RPE > 25) exhibited significantly shorter RPE-time inter-
vals than the left-handers (see Table 3; Mann-Whitney U Test: z =
6.49, p < .001); logarithmic transformation of the RPE-time inter-
vals also yielded the same results (F1, 2385 = 60.66, p < .001). How-
ever, the mean LPE-time interval was significantly shorter in left-
handers than right-handers, for the raw data (z = 2.97, p < .005) and
for the log data (F1, 1063 = 11.22, p = .001).

Time Intervals in Males and Females for RPE and LPE

Mann-Whitney Test indicated that there was a sex difference only
for the RPEs in right-handed rats: the time intervals for the conse-
quent RPEs were significantly shorter in males than females (z =
1686 M. Güven et al.

–5.67, p < .001); the time intervals for the consequent LPEs were
not significantly different in males and females (z = –0.34, p > .70).
In left-handed rats, there was also no significant sex difference for
the consequent time intervals for the LPEs (z = –0.12, p > .90);
however, the male rats had significantly shorter time intervals than
females for the RPEs (z = –2.30, p < .05). The total sample yielded
similar results: the consequent time intervals between the RPEs were
significantly shorter in males than females (z = –6.83, p < .001);
there was, however, no significant sex difference between the con-
sequent time intervals for the LPEs (z = –1.39, p > .15).

DISCUSSION

Global Hand Preference

In accord with the traditional assessment of hand preference, the

numbers of the right- and left-paw entries were counted till the sum
of the paw reaches reached 50. The results of the present work were
consistent with those from the same laboratory (see Elalmis et al.,
2003): there was a population-level right-hand preference in rats
(RHs: 80%, LHs: 13.3%, MHs: 6.7%). In accordance, Pence (2002)
has recently found similar percentages for his 114 Sprague-Dawley
rats: 70.2% RHs, 19.3% LHs, and 11.9% MHs. These results suggest
that the distribution of hand preference in rats is similar to human
hand preference, and reject the uniqueness of men in a population-
level right-handedness (see Elamis et al., 2003 for a review of hand-
edness in animals). The results are also in agreement with previous
studies in rodents (Glick & Ross, 1981; Tang & Verstyenen, 2002;
Tsai & Maurer, 1930; Waters & Denenberg, 1994), and provide
further support for the notion that a homology exists between hand
preference in humans and rats. There are of course other studies
indicating a chance distribution of animal handedness (see for a
review Elalmis et al., 2003).
Considering the strong right-handers, strong left-handers, and in-
between (mixed) handers, the percentages were found to be 35.5%,
4.6%, and 59.9% for the strongly right-handed, strongly left-handed,
and mixed-handed rats, respectively. These proportions are close to
 Paw Preference in Rats 1687

humans: Tan (1988) has reported that the consistent right-handers

comprised 32.4% (35.5% in rats) and mixed-handers comprised 49.5%
(59.9% in rats) in the Turkish university students. Considering the
grasp-reflex asymmetries, the percentages of the strongly right-handed
(35.5%), strongly left-handed (4.6%), and mixed-handed (59.9%)
rats were close to the percentages of the right-hand dominance (25.7%),
left-hand dominance (8.3%), and mixed-handedness (66.0%) in grasp
reflex of human neonates (see Tan & Tan, 1999).
The distribution of the hand preference (RPE scores) in our rats
was not U-shaped opposite to a generally accepted belief (see Bulman-
Fleming, Bryden, & Rogers, 1997). It was, however, J-shaped like
the distribution of hand preference in humans (see Tan, 1988). The
percentages of the right-, left-, and mixed-handedness did not fit a
25:25:50 distribution expected from a binomial chance distribution.
Supporting the above results (i.e., overwhelmingly right-paw pref-
erence in the male and female rats) 99.5% of the right-handed rats,
first used their right paw to reach the food, only 0.5% of these used
the left paw to reach the food. So, the right-handers were really
right-handed in paw preference. Contrary to right-handers, 98.6% of
the left-handed rats first used their left paw to reach the food pallet,
only 1.4% first used their right paw in food reaching. As expected
from a mixed-handed population, nearly half of the mixed-handed
rats (59.0%) first used their right paw, and 41% first used their left
paw in food reaching test. These results indicate that not only the
number of the RPE significantly exceeded the number of LPE in
right-handed rats, the right-handed rats preferred their right hand
and the left-handed rats preferred their left hand for the first food-
reaching task. These results also indicate that a behavioral (psy-
chological) trait (preference) reflects the cerebral motor asymmetry.
The time interval between putting the rats into the test cage and
the first paw reach was different in right-, left-, and mixed-handed
rats: the time for the first RPE was significantly shorter than the
first LPE in right-handed rats, vice versa in left-handed rats; there
was no significant difference between the first RPE and first LPE in
mixed-handed rats. So, the right-handed rats first used their right
paw to reach the food in a shorter time than the left paw; the left-
handed rats first used their left paw in a shorter time than the right
paw; the mixed-handed rats randomly used both paws to reach the
1688 M. Güven et al.

food without any significant difference between times for the first
RPE and LPE for food reaching.
The male rats were faster than the female rats in the first RPE as
well as the first LPE. The male rats were also faster than females in
general food reaching (see RESULTS); the time intervals between
the consequent paw reaches were significantly shorter in males than
females, but the sex difference in the consequent time intervals was
significant only for the RPEs, not for the LPEs in right-handers and
left-handers. This interesting result suggests that the left brain is
important for the speedy timing of the right-paw reaches in right-
and left-handed rats.

Conclusions
The present study introduced a new method to assess handedness in
rats. This was a computerized method to analyze the time intervals
between the consequent paw entries. Contrary to the general belief,
the distributions of the global RPEs (right + left = 50) was J-shaped
not U-shaped. The distribution of the time intervals between the
consequent RPEs was also J-shaped as in human hand preference.
Only after logarithmic transformation of the raw data, did the times
between RPEs and LPEs exhibit a Gaussian (normal) distribution.
The male rats were faster than females; there was a sex difference
in the time intervals for the RPEs, but not for the LPEs, accentuat-
ing the role of the left brain for sex differences in motor control.

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