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Dynamics of Manual Skill: A Computerized Analysis of Single PEG Movements

and Stochastic Resonance Hypothesis of Cerebral Laterality
Derya Deniz Elalmis a; Üner Tan b
a
Medical School, Department of Physiology, Erciyes University, Kayseri, Turkey b Faculty of Sciences,
Department of Physics, Cukurova University, Adana, Turkey

Online Publication Date: 01 March 2008

To cite this Article Elalmis, Derya Deniz and Tan, Üner(2008)'Dynamics of Manual Skill: A Computerized Analysis of Single PEG
Movements and Stochastic Resonance Hypothesis of Cerebral Laterality',International Journal of Neuroscience,118:3,399 — 432
To link to this Article: DOI: 10.1080/00207450701668012
URL: http://dx.doi.org/10.1080/00207450701668012

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 Intern. J. Neuroscience, 118:399–432, 2008
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ISSN: 0020-7454 / 1543-5245 online

DOI: 10.1080/00207450701668012

DYNAMICS OF MANUAL SKILL:

A COMPUTERIZED ANALYSIS OF SINGLE PEG
MOVEMENTS AND STOCHASTIC RESONANCE
HYPOTHESIS OF CEREBRAL LATERALITY
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DERYA DENIZ ELALMIS

Erciyes University
Medical School
Department of Physiology
Kayseri, Turkey

ÜNER TAN
Cukurova University
Faculty of Sciences
Department of Physics
Adana, Turkey

Hand skill was analyzed using a computerized peg moving task. The durations
of single hand movements (PMTs) were accurately measured in right-hand (RH)
and left-hand (LH) writers. One trial consisted of 10 movements of the right
hand and 10 movements of the left hand. Each participant performed five trials.
Women showed significantly higher percentage than men in right-handedness; men
showed higher percentage than women in left-handedness. This sex difference
completely disappeared after taking the same height range in participants. The
mean RH- and LH-PMTs decreased in 5 successive trials, even within a single trial
during 10 successive hand movements, indicating a learning effect of repeated hand
movements. The LH- minus RH-PMTs exhibited fluctuations within a single trial
between positive (faster right hand) and negative (faster left hand) values. LH–RH

Received 28 December 2006.

This study was partly supported by the Turkish Academy of Sciences, Ankara, Turkey.
Address correspondence to Prof. Dr. Uner Tan, Cukurova University, Faculty of Sciences,
Department of Physics, 01330 Adana, Turkey. E-mail: unertan37@yahoo.com

399
 400 D. D. ELALMIS AND U. TAN

PMTs were significantly greater than zero, in favor of right hand, in RH-writers,
but not significantly different from zero in LH-writers, exhibiting a true fluctuating
asymmetry. Participants with no familial sinistrality (FS-) were preponderantly
right-handed (ca. 90%), those with left-handed mother and right-handed father
(FS+1) showed stochastic distribution of hand preference (50:50). Participants
with right-handed mother and left-hander father (FS+2) were not different from
FS- individuals. LH–RH PMT was significantly greater than zero in FS- participants,
almost equal to zero in FS+1 participants, and greater than zero in FS+2 participants
exhibiting greater asymmetry than that in FS- participants. These results suggest a
genetic inheritance of direction and degree of handedness, being a a X-linked trait
originating from mother’s genotype. It was suggested that fluctuating asymmetries
may reflect interactions between stochastic resonance phenomena within right and
left brains. This property of brain may genetically transmitted from mother’s X
chromosome; a net effect of stochastic interactions between hemispheres may
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result in right- or left-handedness, a predominantly unidirectional coupling creating

right-handedness, in favor of left brain, and a stochastic bi-directional coupling
between hemispheres would be a main trait of left-handers. This new “stochastic
resonance hypothesis of cerebral laterality” concerns with stochastic fluctuations
in hand-skill asymmetry and inter-hemispheric coupling through corpus callosum,
and seems to be important for new developments in handedness research.

Keywords cerebral laterality, hand preference, hand skill, peg-moving task, sex,
stochastic resonance

INTRODUCTION
Hand skill is often measured using a peg-moving task (see Annett, 2002,
pp. 332–334). However, this method provides only limited information
regarding the skill of hands, for a number of reasons. First, there are different
time periods between the “start” command, pushing on the button of a hand
chronometer, and initiating contact with the first peg. These time periods may
be different because of varying reaction times of the participants. Second, the
measured peg-moving time (PMT) is the sum of 10 peg movements. This means
that PMT cannot inform one about the hand skill for single hand movements.
Third, the standard deviations of the mean LH–RH PMTs in individual subjects
are not taken into consideration when an average of 10 hand movements is
calculated in constructing the histograms for the LH–RH PMTs. The right-shift
theory is based entirely on such a global analysis of hand skill (see Annett, 2002,
pp. 48–72).
The authors thought that, instead of measuring the total PMT using a
push-button hand chronometer, they could electronically measure every single
PMT without the investigator’s intervention. In doing so, the study could use
 HAND SKILL 401

every single hand movement in statistical analysis rather than using simple
means from single subjects without using their SDs. In addition, the study
could obtain some information about the dynamics of hand skill, reflecting
brain dynamics at the same time.
The primary aim of this work was to study the changes in the single
RH-, LH-, and LH minus RH PMTs, using a computerized peg-moving task,
which could measure PMTs in ms accuracy for each hand movement. In fact,
the peg-moving task was computerized by Curt, De Agostini, Maccario, and
Dellatolas in 1995. De Agostini and Dellatolas (2001) also used this apparatus
in subsequent work performed with normal children. They measured the mean
PMT for 10 hand movements without analysing the single PMTs.
The present authors proposed that changes in single PMTs would help
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one understand the brain’s dynamic role in movement timing and learning
through the repetition of hand movements. Learning effects from repetitive
hand movements were expected, as repetition is the essence of motor learning.
Learning in peg-moving tasks has been first demonstrated by Tan (1989).
Furthermore, the brain is a dynamic system, that is, movement control may
change over time. This cannot be measured with a simple hand chronometer
measuring the total PMT of 10 hand movements. Accordingly, the authors
computerized the peg-moving task to provide measurements accurate enough
to understand the brain dynamics of bimanual movement.
To obtain more and accurate information about the dynamics of hand skill
in conjunction with brain dynamics, the study measured single PMTs from right
and left hands, using a computerized peg-moving task. Within the framework
of this analysis, the study compared data from inter- and intra-subject analysis;
the authors looked for stochastic dynamics of hand skill following stochastic-
resonance properties of neural systems; the authors asked whether repeated
hand movements in peg-moving task would result in motor learning within
a single trial and following five trials, and if the dynamic properties of hand
skill genetically determined. Accordingly, a new hypothesis was created about
the origins of handedness with respect to stochastic resonance phenomena in
interacting neural systems.

METHOD
The participants were 196 university students (90 men and 106 women, 18–24
years) from the medical school of Cukurova University (Adana, Turkey). They
were without psychological or neurological signs and symptoms, and were
randomly recruited. Hand preference was determined according to writing
 402 D. D. ELALMIS AND U. TAN

hand, that is, right-handers (n = 175) always used the right hand and left-
handers (n = 21) the left hand to write legibly. Handedness can indeed be
divided into RH-writers and LH-writers, because writing hand is useful for
a scheme of classification according to an association analysis (see Annett,
2002, p. 37). Therefore, RH-writers and LH-writers were considered as separate
categories in the present work.
Manual skill was determined using the peg-moving task (see Annett, 2002,
pp. 332–334) with a wooden board of the same dimensions as Annett’s. There
were 2 parallel rows 20.3 cm apart, each comprising 10 holes 2.5 cm apart.
Participants stood in front of this pegboard, facing down the rows. The single
trial consisted of placing pegs from a hole in one row to the adjacent hole in
the opposite row, without dropping a peg. Participants started with their right
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hands, moving the 10 pegs from right to left rows; participants then used their
left hands to move the 10 pegs from left to right rows, giving a total of 20 peg
movements for the trial. The subjects repeated the same procedure five times
(five trials).
The durations of single peg movements were measured using a comput-
erized version of Annett’s original apparatus. For this purpose, sensors were
placed at the bottoms of the holes and connected over an electronic circuit to
the parallel port of a computer. The duration between the start and the end
of each single peg-movement was recorded using computer software written
in C++ and run on the Windows 98 operating system; the set-up included
a timer that sampled at 1 ms intervals. Participants were instructed to move
the pegs from one row to the other as quickly as possible after the start of
the trial. The software automatically stopped timing when 10 pegs had been
moved from one row to the other. At the end of the experiment, the data were
saved on the hard disk and statistically analyzed using SPSS for Windows (V.
11.0). Effect size (η2 ) was estimated giving a partial eta-squared value, which
described the proportion of total variability attributable to a factor. The PMTs
for the first hand movements were discarded from the statistical analysis, as
their movement times were half as long of the consequent hand movements.

RESULTS
Inter-Subject Analysis
Hand Preference; Sex Differences. The numbers and percentages of the
male and female RH- and LH-writers are presented in Table 1. The numbers
of the RH-writing women (n = 99, 93.4%) significantly exceeded the number
 HAND SKILL 403

Table 1. Height and weight in right- and left-handed men and women

Weight (Kg) Height (cm)

Participants n Mean SEM Mean SEM

RH-writers 175 63.1 0.89 168.7 0.69

Men 76 72.5 1.18 176.2 0.74
Women 99 55.8 0.68 162.9 0.60
LH-writers 22 69.5 2.38 174.0 2.02
Men 15 76.2 1.35 178.4 2.02
Women 7 55.1 1.60 164.6 1.60
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of the RH-writing men (n = 76, 83.5%), and vice versa in LH-writers, that is,
significantly more LH-writing men (n = 15, 16.5%) than LH-writing women,
n = 7, 6.6%, (χ 2 = 4.82, p < .05). In the total sample, there were 175 RH-writers
(88.2%) and 22 LH-writers (11.2%).
Because of hormonal influences on both handedness and bodily measures,
the mean weights and heights of the right- and left-handed men and women were
calculated (see Table 1). In RH- and LH-writers, handedness was a significant
factor for height, F(1, 196), p < .05, η2 = .033, and weight, F(1,196) =
5.89, p < .06, η2 = .029. Namely, the LH-writers were heavier and taller than
the RH-writers. This was actually expected because there were significantly
more LH-men than LH-women; men were significantly taller and heavier than
women. Although the mean weight and height were greater in consistently
right-handed men than in consistently right-handed women, the difference was
statistically not significant, F(1, 176) = 2.63, p > .10 for height, and F(1,176) =
2.61, p > .10 for weight.
Following the aforementioned results, the sex difference in hand preference
was re-investigated, taking the body size (height) into consideration. In RH- and
LH-writers, the heights ranging from 166.0 to 172.0 cm was taken as a common
range for men (range = 163 to 193 cm) and women (range = 152 to178 cm).
In this height range (166.0–172.0 cm), the sex difference in hand preference
completely disappeared: 88.0% (n = 24) of women and 90.0% (n = 29) of men
were RH-writers, 5 women (12.0%) and 5 men (10.0%) were LH-writers, the
differences being statistically not significant (χ 2 = 0.56, p > .55).
The same was also true for weight, which ranged from 52.0 to 93.0 Kg
in men and from 44.0 to 74.0 Kg in women. As a common weight ranging
from 54.0 to 72.0 Kg was taken into consideration, the sex difference in hand
preference for writing completely disappeared (χ 2 = 0.95, p > .60): 93.0%
 404 D. D. ELALMIS AND U. TAN

Table 2. RH-, LH-, and LH- minus RH-PMTs in RH-writers and LH-writers

RH-PMT LH-PMT LH–RH PMT

Subjects n Mean SE Mean SE Mean SE

Total (1st trial) 197 1.465 0.012 1.540 0.011 0.075 0.010
Men 91 1.483 0.019 1.535 0.016 0.053 0.014
Women 106 1.450 0.016 1.543 0.015 0.093 0.014
Total (5th trial) 197 1.317 0.012 1.437 0.013 0.123 0.011
Men 91 1.310 0.020 1.409 0.020 0.106 0.015
Women 106 1.324 0.014 1.461 0.018 0.137 0.015
RH-writers
1st trial 175 1.455 0.013 1.539 0.012 0.084 0.011
Men 76 1.466 0.020 1.527 0.018 0.062 0.016
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Women 99 1.446 0.017 1.548 0.016 0.101 0.015

5th trial 175 1.306 0.012 1.449 0.014 0.143 0.010
Men 76 1.288 0.020 1.426 0.021 0.138 0.014
Women 99 1.321 0.014 1.467 0.019 0.146 0.015
LH-writers
1st trial 22 1.551 0.034 1.541 0.031 −0.001 0.022
Men 15 1.573 0.047 1.576 0.039 0.010 0.030
Women 7 1.503 0.026 1.467 0.039 −0.026 0.031
5th trial 22 1.403 0.047 1.339 0.041 −0.037 0.028
Men 15 1.421 0.060 1.324 0.057 −0.058 0.032
Women 7 1.365 0.077 1.372 0.050 0.007 0.055

of men (n = 40) was RH-writers and 7% (n = 3) was LH-writers; 93.3% (n =

56) of women was RH-writers and 6.7% (n = 4) LH-writers.

Mean PMTs in Single Participants. The means with their SEMs of PMTs
for the first and fifth trials are presented in Table 2. For each participant, a mean
PMT was calculated using nine RH- and nine LH-movements, that is, each
subject was represented by a single mean. Namely, the SEMs did not belong
to the mean PMTs for single individuals, as is in the traditional analysis of
peg-moving task.
As the sex and handedness were taken as between-subjects factors, and
RH-, LH-, and LH–RH PMTs as dependent variables, first trial, it was found
for the RH-PMT that sex was not significant (F = 1.22, p > .25), whereas
handedness was significant (F = 4.13, p < .05): the mean RH-PMT was
greater in LH-writers than in RH-writers. For the LH-PMT in the first trial, sex
and handedness were not significant factors (F = 1.38, p > .20 for sex; F =
 HAND SKILL 405

0.20, p > .65 for handedness). For the LH–RH PMT, sex was not significant
(F = .00, p > .95), but handedness was significant (F = 7.45, p < .01): the
mean LH-RH PMT was greater in RH-writers than in LH-writers.
In the fifth trial, sex was not a significant factor for the RH-PMT (F =
0.09, p > .75), while handedness was significant (F = 5.21, p < .05): the mean
RH-PMT in LH-writers was greater than that in RH-writers. For the LH-PMT,
sex again was not a significant factor (F = 0.97, p > .30), but handedness was
significant (F = 4.79, p < .05): the mean LH-PMT was greater in RH-writers
than that in LH-writers. For the LH–RH PMT, sex was not significant (F =
1.02, p > .80), whereas handedness was significant (F = 26.34, p < .001,
η2 = .121): the mean LH–RH PMT was greater in RH-writers than that in
LH-writers, in favor of the right hand.
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In the total sample (n = 197), first trial, the mean LH-PMT was significantly
greater than the RH-PMT (MD = 0.075; t = 4.58, df = 392, p < .0001). The
LH–RH PMT was significantly greater than zero (MD = 0.075 s; t = 7.54,
df = 196, p < .0001). In the fifth trial, the mean LH-PMT was significantly
greater than the mean RH-PMT (MD = 0.120 s; t = 6.73, df = 392, p < .0001).
The mean LH–RH PMT was significantly greater than zero (MD = 0.123; t =
11.67, df = 196, p < .0001). The mean RH-PMT was significantly smaller in
the fifth than the first trial (MD = 0.148 s, t = 8.79, df = 392, p < .0001), as
was for the LH-PMT (MD = 0.103 s; t = 6.53, df = 392, p < .0001). The mean
LH–RH PMT in the first trial was significantly greater than that in the fifth trial
(MD = 0.048 s; t = 3.32, df = 392, p = .001). There was no significant sex
difference in the mean RH- and LH-PMTs in the first trial, F = 2.17, p > .10,
while the mean LH- minus RH PMT was significantly greater in women than
in men, F(1, 196) = 4.10, p < .05. In the fifth trail, there was no significant
sex difference in the mean RH-, LH-, and LH-RH-PMTs (F = 2.00, p >
.10).
In RH-writers (n = 175), first trial, sex was not a significant factor for the
mean RH- and LH-PMTs (F = 0.58, p > .60). The mean LH–RH PMT was
greater in women than in men, the difference being only marginally significant
(F = 3.51, p = .06). This small sex difference disappeared, however, after
taking the height of the participants as a covariate (F = 1.97, p > .15). The
mean LH-PMT was significantly greater than the mean RH-PMT; the LH–RH
PMT was significantly greater than zero (t = 7.91, df = 174, p < .001). In
the fifth trial, there was no significant sex difference in the RH- and LH-, and
LH–RH PMTs (F = 1.14, p > .25). After taking the height as a covariate, the
mean LH–RH PMTs were found to be equal in men and women (F = .000, p >
.99). The mean LH-PMT was significantly greater than the mean RH-PMT; the
 406 D. D. ELALMIS AND U. TAN
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Figure 1. Distributions of LH-RH PMTs (abscissa) in RH-writers (A, B) and LH-writers (C, D)
for first (A, C) and fifth (B, D) trials.

mean LH–RH PMT was significantly greater than zero (t = 13.78, df = 174,
p < .001).
In LH-writers (n = 22), first trial, there was no significant difference
between the mean RH- and LH-PMTs; the mean LH–RH PMT was not
significantly different from zero (t = 0.06, df = 21, p > .90). Sex was not
a significant factor for the RH-, LH-, and LH–RH PMTs (F = 1.24, p > .20).
In the fifth trial, there was no significant sex differences in the RH-, LH-, and
LH–RH PMTs (F = 0.30, 0.28, and 1.14, p > .55, .60, and .25, respectively).
There was no significant difference between the mean RH- and LH-PMTs; the
mean LH–RH PMT was not significantly different from zero (t = 1.33, df =
21, p > .15).

Distributions of LH–RH PMTs. Figure 1 illustrates the distributions of the

LH–RH PMTs for the first (A, C) and fifth trials (B, D) in RH-writers (A, B)
 HAND SKILL 407

and LH-writers (C, D). All of them fitted to Gaussian distributions: for A, KS
z = 0.76, p > .60; for B, z = 0.94, p > .30; for C, z = 0.83, p > .50; for D, z =
0.92, p > .37.
In RH-writers (n = 175) , 28.6% (n = 50) of the participants had a faster
left hand, and 71.4% (n = 125) had a faster right hand during the first trial. The
proportion of the RH-writers with a faster left hand decreased to 13.7% (n =
24) and that with a faster right hand increased to 86.3% (n = 151) within the
fifth trial, the difference being statistically significant (χ 2 = 10.71, df = 1, p =
.001).
Of 22 LH-writers, 10 participants (45.5%) had a faster left hand and 12
participants (55.5%) had a faster right hand in the first trial, the difference
between these proportions being statistically not significant (χ 2 = 0.09, df =
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1, p > .75). These proportions did not change in the fifth trial (see Figure 2D).

Relations of LH-Minus RH-PMTs to RH- and LH-PMTs. The relations

of the LH–RH PMTs to the RH- and LH-PMTs are illustrated in Figure 2.
In RH-writers (n = 175), there was a negative linear correlation between the
LH–RH PMT and RH-PMT (A: open circles, dashed line, r = –.50, t = 7.55,
p < .001). A similar relationship was found in LH-writers (n = 22), A: dots,
straight line. r = –.55, t = 2.95, p < .01). There was a positive correlation
between the LH–RH PMT and the LH-PMT in RH-writers (B: open circles,
dashed line, r = .37, t = 5.19, p < .01), but there was no significant correlation
between these variables in LH-writers (B: dots, straight line, r = .19, t = 0.88,
p > .35). These results were due to the first trial.
In the fifth trial of RH-writers, the LH–RH PMT inversely correlated with
the RH-PMT (C: open circles, dashed line, r = –.19, t = 2.50, p < .05), but
with a less significant correlation compared to the first trial. In LH-writers,
there was a similar correlation between these variables (C: dots, straight line,
r = –.61, t = 3.46, p < .005). The LH-PMT was found to be significantly
correlated with the LH–RH PMT in RH-writers (D: open circle, dashed line,
r = .58, t = 9.46, p < .001), but no significant correlation was found between
these variables in LH-writers (D: dots, straight line, r = .21, t = 0.95, p > .35).

Intra-Subjects Analysis
The mean RH-, LH, and LH–RH PMTs for the male and female RH- and
LH-writers are presented in Table 3. In this section, the PMTs for single hand
movements were taken into consideration. In doing so, the deviations in single
 408 D. D. ELALMIS AND U. TAN
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Figure 2. Distributions of LH–RH PMTs in RH- and LH-writers. A: RH-writers, first trial; B:
RH-writers, fifth trial; C: LH-writers, first trial; D: LH-writers, fifth trial.

PMTs during a single trial could be analyzed instead of the mean PMTs for
single subjects.

RH-, LH-, and LH–RH PMTs. In the first trial, as the sex and handedness
were taken as between-subjects factors and the RH-, LH-, LH–RH PMTs as the
dependent variables, it was found for the RH-PMT that sex was not significant
(F = 1.92, p > .15), but handedness was significant (F = 16.25, p < .001,
η2 = .010): the mean RH-PMT was significantly greater in LH-writers than
in RH-writers. As the height was taken as a covariate, sex was found to be
a significant factor for the RH-PMT (F = 7.45, p < .01, η2 = .004): the
mean RH-PMT was significantly greater in men than that in women. For the
 HAND SKILL 409

Table 3. Means and SEMs for single PMTs in RH- and LH-writers

Subjects n Hands RH-PMT SEM LH-PMT SEM LH–RH PMT SEM

Total (1st)
Men 197 1773 1.451 0.006 1.592 0.006 0.142 0.006
Women 91 819 1.465 0.009 1.591 0.009 0.127 0.009
Total 106 954 1.439 0.008 1.594 0.008 0.156 0.009
(5th) 197 1773 1.314 0.006 1.437 0.006 0.122 0.007
Men 91 819 1.304 0.009 1.409 0.009 0.105 0.010
Women 106 954 1.323 0.007 1.460 0.008 0.138 0.009
RH-wri.
1st trial 175 1575 1.440 0.007 1.601 0.007 0.161 0.007
Men 76 684 1.449 0.010 1.599 0.010 0.151 0.010
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Wom 99 891 1.434 0.009 1.602 0.009 0.169 0.009

5th trial 175 1575 1.305 0.006 1.449 0.006 0.144 0.007
Men 76 684 1.287 0.010 1.426 0.010 0.139 0.011
Wom 99 891 1.320 0.008 1.467 0.009 0.147 0.010
LH-wri.
1st trial 22 198 1.532 0.017 1.527 0.018 −0.001 0.016
Men 15 135 1.546 0.023 1.550 0.023 0.011 0.019
Wom 7 63 1.503 0.022 1.478 0.023 −0.026 0.031
5th trial 22 198 1.384 0.019 1.339 0.018 −0.043 0.020
Men 15 135 1.398 0.025 1.324 0.023 −0.067 0.025
Wom 7 63 1.365 0.028 1.371 0.026 0.007 0.030

LH-PMT, sex was not significant (F = 2.72, p > .05), but handedness was
significant (F = 17.80, p < .001, η2 = .010): the mean LH-PMT was greater in
RH-writers than that in LH-writers. As the height was taken as a covariate, sex
was found to be a significant factor (F = 3.91, p < .05, η2 = .002): the mean
LH-PMT was greater in men than in women. For the LH–RH PMT, sex was
not significant (F = 0.18, p > .65), handedness was significant (F = 61.20,
p < .001, η2 = .034): the mean LH–RH PMT was greater in RH-writers than
in LH-writers. Height as a covariate did not change these results.
In the fifth trial, for the RH-PMT, sex was not significant (F = 0.02, p >
.85), but marginally significant after taking the height as a covariate (F = 2.58,
p = .07): the mean RH-PMT was greater in men than in women. Handedness
was significant (F = 15.37, p < .001, η2 = .009): the mean RH-PMT was
greater in LH-writers than in RH-writers. For the LH-PMT, sex was significant
(F = 4.66, p < .05, η2 = .009): the mean LH-PMT was greater in men than
in women, but there was not a significant sex effect after taking height as a
covariate (F = 0.17, p > .65). Handedness was a significant factor for the
 410 D. D. ELALMIS AND U. TAN

LH-PMT (F = 22.97, p < .001, η2 = .013): the mean LH-PMT was greater in
RH-writers than in LH-writers. For the LH–RH PMT, sex was not significant
(F = 3.18, p > .05), but handedness was significant (F = 58.18, p < .001, η2 =
.032). This result was not changed under covariance of the height.
In the total sample (n = 197), there were 1773 right- and 1773 left-hand
movements in one trial. In the first and fifth trials, the mean LH-PMT was
significantly greater than the mean RH-PMT (1st trial: t = 16.24, df = 3544,
p < .0001; 5th trial: t = 14.33, df = 3544, p < .0001). The mean RH- and
LH-PMTs in the first trial were significantly greater than those in the fifth trial
(t = 16.28, df = 3544, p < .0001 for RH-PMTs; t = 17.85, df = 3544, p <
.0001 for LH-PMT). In the first and fifth trials, there were no significant sex
difference in the RH- and LH-PMTs (1st trial: F = 1.61, p > .20; 5th trial:
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F = 2.58, p > .05). The mean RH-PMT was significantly greater in LH-writers
than in RH-writers, and vice versa for the LH-PMT (F = 32.53, p < .001, η2 =
.035). Sex∗ handedness was not significant (F = 1.67, p > .15).
The mean LH–RH PMT was found to be significantly smaller in the fifth
trial than in the first trial (t = 2.15, df = 3544, p < .05). The mean LH–RH PMT
in the first and fifth trials showed significant sex differences: it was significantly
greater in women than in men in the 1st trial, F(1, 1772) = 5.25, p < .001, and
in the fifth trial, F(1, 1772) = 5.73, p < .001.
In the first trial, the mean LH–RH PMT was significantly greater than zero
in RH-writers (MD = 0.161 ± 0.007 s, t = 23.89, df = 1574, p < .0001),
whereas it was not significantly different from zero in LH-writers (MD =
–0.001 ± 0.226 s, t = 0.08, df = 188, p > .90). In the fifth trial, the mean
LH–RH PMT was significantly greater than zero in RH-writers (MD = 0.144
± 0.008, t = 20.28, df = 1574, p < .0001), whereas it was significantly smaller
than zero in LH-writers (MD = −0.043, t = 2.17, df = 188, p < .05). The
mean LH–RH PMTs were not significantly different in the first and fifth trials
of the RH men (t = 0.81, df = 1366, p > .40) and women (t = 1.69, df =
1780, p > .05), although the raw data showed smaller values in the fifth than
the first trials. In LH-writing men, the positive mean LH–RH PMT in the first
trial (Mean = 0.011 ± 0.216) reversed to a negative mean (–0.067 ± 0.28) in
the fifth trial, the difference being statistically significant (t = 2.48, df = 250,
p = .01). There was not a significant change in the mean LH–RH PMT in the
first and fifth trials in LH-writing women (t = 0.007, df = 124, p > .40).

Distribution of LH–RH PMT in RH-Writers and LH-Writers. The distribu-

tions of the LH–RH PMTs in RH-writers (A, B) and LH-writers (C, D) for the
 HAND SKILL 411

first (A, C) and fıfth (B, D) trials are shown in Figure 3. In RH-writers, the first
trial of 1575 hand pairs, 54 (3.4%) LH–RH PMTs were equal to zero, 1124
(71.4%) were greater than zero (faster right hand), and 397 (25.2%) less than
zero (faster left hand). For the fifth trial (see Figure 2B) of RH-writers, 1089
(69.1%) were greater than zero (faster right hand), 426 (27.0%) were less than
zero (faster left hand), and 60 (3.8%) were equal to zero (tied). The proportions
in the first and total trials were not different in RH-writers (χ 2 = 1.89, p >
.15).
In the first trial of LH-writers (n = 22, 198 hand pairs; Figure 2C), 4.5%
(nine hand pairs) of the participants had equal LH–RH PMTs, 41.4% (82 hand
pairs) had a faster right hand, and 54.0% (98 hand pairs) a faster left hand.
The difference between the LH–RH PMTs with faster right and left hands
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was statistically significant (χ 2 = 6.10, df = 1, p < .05). For the fifth trial
(Figure 3D), of 198 LH–RH PMTs, 8 (4.0%) were equal, 77 were greater
than zero (38.9%; faster right hand) and 113 (57.1%) less than zero (faster
left hand). In this sample, the number of the LH–RH PMTs with a faster left
hand significantly exceeded that with a faster right hand (χ 2 = 62.62, df =
1, p = < .0001). There was no significant difference between the proportions
of the participants with faster left hand in the first and fifth trials (χ 2 = 0.27,
p > .60).

Effects of Repetition on PMT

Mean PMTs in Consequent Five Trials. As repetition is the essence of
learning, the RH- and LH-PMTs in the consequent five trials were subjected to
a statistical analysis. Figure 3 illustrates the changes in the mean PMTs during
the consequent five trials (abscissa) for the RH-PMTs (A), and LH-PMTs (B)
in RH-writers (circles) and LH-writers (triangles). In RH-writers (n = 175), the
mean RH-PMTs (A: circles) and LH-PMTs (B: circles) significantly decreased

Table 4. Mean RH-, LH-, and LH-RH PMTs (s) in FS-, FS+1, and FS+2 participants

Right hand Left hand LH-RH

Subjects n Hands Mean SE Mean SE Mean SE

FS- 150 2700 1.43 0.005 1.55 0.005 0.12 0.005

FS+1 6 108 1.46 0.025 1.50 0.025 0.04 0.024
FS+2 11 198 1.30 0.015 1.54 0.017 0.24 0.016
 412 D. D. ELALMIS AND U. TAN
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Figure 3. RH- (A) and LH-PMTs (B) (means ± 2.0 SEMs) in RH-writers (open circles) and
LH-writers (triangles) for the five consequent trials (abscissa).

after the consequent five trials: F(4) = 83.73, p < .001 for the RH-PMTs; F(4) =
98.78, p < .001 for the LH-PMTs, with significant quadratic trends (F = 47.16,
p < .001 for RH-PMTs and F = 54.00, p < .001 for the LH-PMTs).
In LH-writers (n = 21), the differences between the mean PMTs during
the five consequent trials were found to be significant for the mean RH-PMTs
(F(4) = 17.30, p < .001; A: triangles) and for the LH-PMTs (F(4) = 22.96, p <
.001; B: triangles). The trends were significantly quadratic for the RH-PMTs
(F = 8.61, p < .005) and for the LH-PMTs (F = 21.05, p < .001).
 HAND SKILL 413

In contrast to RH- and LH-PMTs, LH–RH PMTs did not show a significant
trend during the five trials in RH-writers and LH-writers (see Figure 6C): F =
0.81, p > .45 for RH-writers; F = 1.84, p > .10 for LH-writers.

Single PMTs During a Single Trial. To examine the learning effect of

repeated hand movements during one single trial, the changes in the single
PMTs of the consequent nine hand movements were examined. Figure 4
shows the mean PMTs for single hand movements, but only for the first trial,
which consisted of nine single hand movements. In RH-writers, the means
of the consequent RH- (A: circles, straight lines) and LH-PMTs (B: circles,
straight lines) appeared to decrease from the first to the last hand movement;
the differences between the mean PMTs of the consequent hand movements
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(trends) were found to be statistically significant: F(8, 1566) = 15.62, p <

.001 for the RH-PMTs; F(8, 1566) = 15.66, p < .001 for the LH-PMT, but the
mean LH–RH PMTs did not show significant changes with consequent hand
movements: F(8,1566) = 0.77, p > .60 (C: circles, straight lines). In LH-writers
(triangles, dashed lines), the differences between the mean consequent RH- and
LH-PMTs were not significant: F(8, 180) = 0.84, p > .55 for the RH-PMTs
(A: triangles, dashed lines); F(8,180) = 0.99, p > .40 for the LH-PMTs (B:
triangles, dashed lines); and F(8, 180) = 0.79, p > .60 for the LH–RH PMTs
(C: triangles, dashed lines).

Fluctuations in Single LH–RH PMTs. The single LH–RH PMTs usually

fluctuated between positive (a faster right hand) and negative (a faster left
hand) values during nine single hand movements in RH- and LH-writers. The
mean LH–RH PMTs during the first trial (squares) with their ± 2.0 SEMs
(vertical bars) for the RH-writers (A) and LH-writers (B) are shown in Figure
5. With regard to the between-subject variabilities (fluctuating asymmetry
between subjects in hand skill), of the 175 RH-writers, 25 (14.3%) had negative
means (faster left hand), and 150 (85.7%) had positive means (faster left hand)
That is, not all of the right-handed subjects had a faster right hand; 14.3% of
them exhibited a faster left hand. With regard to the single peg movements in
RH-writers, the sign-test showed that of the 1575 hand pairs, 1,123 (71.3%)
LH–RH PMTs were greater than zero (faster right hand), 398 (25.3%) LH–RH
PMTs were less than zero (faster left hand), and 54 (3.4%) were equal to zero
(tied).
Of the 22 LH-writers, 9 (42.9%) had negative means (LH–RH < 0),
exhibiting a faster left hand, and 12 (67.1%) had positive means (LH–RH >
0), exhibiting a faster right hand in the first trial. The difference between the
 414 D. D. ELALMIS AND U. TAN
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Figure 4. Means and SEMs for the RH- (A), LH- (B), and LH–RH (C) PMTs during the consequent
nine hand movements in the first trial.
 HAND SKILL 415
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Figure 5. Mean LH–RH PMTs with ± 2.0 SEMs (ordinate) in single subjects (abscissa) during
first trial. The numbers refer to the subjects’ identifications. Above: RH-writers; below: LH-writers.

numbers of the subjects with faster right and left hands was not significant
(χ 2 = 0.19, p > .65). With regard to the single hand movements (189 LH–RH
PMTs), 82 (43.4%) exhibited a faster right hand (LH–RH > 0), 98 (51.9%)
a faster left hand (LH–RH < 0), and 9 (4.8%) LH–RH PMTs were equal to
zero. There was no significant difference between the numbers of the LH–RH
PMTs being greater and less than zero: χ 2 = 1.25, p > .25. The numbers of the
positive and negative signs were not significantly different (z = 1.12, p > .25).
The number of mismatched LH–RH PMTs, that is, negatives in right-handers
and positives in left-handers, was significantly greater in left-handers than in
right-handers (χ 2 = 58.74, p < .001).

Fluctuations During the First and Last Halves in a Single Trial. To answer
the question of whether there may be differences in the fluctuations with time,
the numbers of positive and negative LH–RH PMTs were counted in the first
 416 D. D. ELALMIS AND U. TAN

half (five LH–RH PMTs) and in the last half (four LH–RH PMTs) of the first
trial in RH-writers and LH-writers, and the numbers of the positive and negative
signs were compared using the sign test. There was no significant difference
between the numbers of the positive, negative, and equal signs of the first and
last halves either in RH-writers (χ 2 = 1.19, df = 2, p > .55) or in LH-writers
(χ 2 = 1.89, df = 2, p > .35).

Fluctuations During First and Last Trials. In RH-writers (n = 175), during

the first trial, 1,123 of the 1,575 LH–RH PMTs (71.3%) were greater than zero
(faster right hand), 398 (25.3%) less than zero (faster left hand), and 54 (3.4%)
equal to zero (tied). During the last trial, 1089 (69.1%) were greater than zero,
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426 (2.0%) less than zero, and 60 (3.8%) equal to zero (tied). There was no
significant difference between these proportions in the first and last trials (χ 2 =
1.89, df = 2, p > .35). In LH-writers (n = 22), during the first trial, 82 of the 189
LH–RH PMTs (41.4%) were greater than zero (faster right hand), 107 (54.0%)
were less than zero (faster left hand), and 9 (4.6%) were equal to zero. In the
fifth trial, of 198 LH-RH PMTs, (38.9%) were greater than zero (faster right
hand), 113 (57.1%) less than zero (faster left hand, and 8 (4.0%) equal to zero.
The difference between the numbers of the faster left hand in the first and fifth
trials was statistically not significant (χ 2 = 0.16, p > .65). The proportion of
the mismatched negativities (25.3%) in RH-writers significantly exceeded the
proportion of the mismatched positivities (41.4%) in LH-writers (χ 2 = 22.27,
p < .0001). The proportion of the matched positivities (71.3%) in RH-writers
significantly exceeded the matched negativities (54.0%) in LH-writers (χ 2 =
23.97, p < .0001). There was no significant difference between the equal RH-
and LH-PMTs in RH-writers and LH-writers (χ 2 = 0.43, p > .50).

Fluctuations in RH- and LH-PMTs. To understand the fluctuations in single

hand movements of right- and left-handed men and women, the standard
deviations of the mean PMTs as a measure of dispersion were compared.
In the total sample, the left-hand PMT showed significantly more fluctuations
than the right-hand PMT in women (F = 1.27, p < .005) and men (F = 7.84,
p < .001). In RH- and LH-writers, there were significantly more fluctuations
in the left-hand PMT than the RH-PMT (F = 1.20, p = .005 for right-handers;
F = 1.61, p < .05 for left-handers). In contrast to the single right- and left-hand
movements, the fluctuations (three times standard deviations) in the LH–RH
PMTs did not show significant differences between men and women (F = 1.01,
p > .90), and between RH- and LH-writers (F = 1.01, p > .95).
 HAND SKILL 417

Familial Sinistrality
Hand Skill. To have an idea about a possible role of genetics in hand-skill, the
distributions of the RH-, LH-, and LH–RH PMTs were analyzed in participants
with no familial sinistrality (FS-), with left-handed mother–right-handed
father (FS+1), and with left-handed father–right-handed mother (FS+2). The
distributions of the LH-RH PMTs in FS-, FS+1, and FS+2 participants are
illustrated in Figure 6 (first two trials). The distributions were normal in FS-
(KS z = 1.22, p > .10), FS+1 (KS z = 0.67, p > .75), and FS+2 (KS z = 0.93,
p > .30) individuals. In FS-individuals, the mean LH–RH PMT was found to
be significantly greater than zero (t = 23.26, df = 2699, p < .001). The FS+1
individuals (n = 5, hands = 90) entirely consisted of males; the mean LH–RH
PMT was not significantly different from zero in these individuals (t = 1.54,
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df = 89, p > .10). In FS+2 individuals, the mean LH–RH PMT was found to
be significantly greater than zero (t = 14.75, df = 197, p < .001).
The mean LH–RH PMT was significantly smaller in FS+1 than in FS-
individuals (MD = 0.08 s, t = 2.68, df = 2788, p < .01); it was significantly
greater in FS+2 individuals than in FS+1 individuals (MD = 0.20 s, t =
6.84, df = 286, p < .0001); it was significantly greater in FS+2 than in FS-
individuals (MD = 0.12 s, t = 5.89, df = 2896, p < .0001).
In FS-participants, 66.4% exhibited positive LH–RH PMTs (faster right
hand), 30.6% negative LH–RH PMTs (faster left hand), and 3.0% ties (LH–RH
PMT = 0). The difference between these proportions was found to be
statistically significant (sign test, z = 18.90, p < .0001). In FS+1 individuals,
52.2% had positive signs, 44.4% negative signs, and 3.3% ties, the difference
being statistically not significant (z = 0.64, p > . 50). In FS+2 individuals,
85.4% had positive signs, 10.6% negative signs, and 8.9% ties, the differences
being statistically significant (z = 10.67, p < .0001). The percentage of
positive signs decreased from 66.4% in FS- participants to 52.2% in FS+1
individuals, the difference (14.2%) being statistically significant (χ 2 = 7.2,
p < .01). Contrarily, the percentage of the positive signs increased from 66.4%
in FS- individuals to 85.4% in FS+2 individuals, the difference (19.0%) being
significant (χ 2 = 20.6, 7.2, p < .0001). On the other hand, the percentage of
the negative signs (faster left hand) significantly increased from 30.6% in FS-
individuals to 44.4% in FS+1 individuals (χ 2 = 7.1, p < .01), and significantly
decreased to 10.6% in FS+2 individuals (χ 2 = 34.7, p < .0001).
There was no significant difference between the mean RH-PMTs of the
FS- and FS+1 individuals (MD = 0.05 s, t = 1.27, df = 1393, p > .20); it
was significantly smaller in FS+2 than FS+1 individuals (MD = 0.18 s, t =
 418 D. D. ELALMIS AND U. TAN
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Figure 6. Distributions of LH-RH PMTs in FS- (A), FS+1 (B), and FS+2 (C) individuals.
Abscissa: LH-RH PMT (s); ordinate: number of hand pairs.
 HAND SKILL 419

4.62, df = 142, p < .0001); it was also significantly smaller in FS+2 than
FS- individuals (MF = 0.13, t = 4.87, df = 1447, p < .0001). Concerning the
mean LH-PMTs, the mean LH-PMT was significantly smaller in FS+1 than
in FS- individuals (MD = 0.05, t = 2.02, df = 2806, p < .05). There were
no significant differences between FS- and FS+2 individuals (MD = 0.01, t =
0.36, df = 1447, p > .70), and FS+1 and FS+2 individuals (MD = 0.04, t =
0.77, df = 142, p > .40).

Hand Preference. Of 150 FS-participants, 136 (90.7%) were RH-writers (81

women and 55 men) and 14 (9.3%) were LH-writers (7 women and 7 men).
There were 6 FS+1 participants, who entirely consisted of men, 50% being
RH-writers and 50% being LH-writers. Of 11 FS+2 participants, 10 (90.9%)
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were RH-writers (4 women and 6 men) and only one (men) was LH-writer
(9.1%).

DISCUSSION
This work presented new results about hand skill measured by the authors’
newly developed computerized analysis of peg-moving task. The traditional
analysis dealt with the mean PMTs for single individuals. This neglected the
SDs of the mean PMTs (10 hand movements in 5 trials) for single participants.
Moreover, the genuine differences in the reaction times of different participants
and different researchers may be the additional disadvantages for the hand–eye
driven peg movements. The study analyzed, instead, the times for single hand
movements in ms accuracy without the researcher‘s intervention.

Sex Differences
Hand Preference. The relative number of right-handed women significantly
exceeded the relative number of right-handed men; the number of left-handed
men significantly exceeded that of left-handed women. This is consistent with
the generally accepted notion about the sex difference in hand preference
reported universally in countries such as in England (Oldfield, 1971), USA
(Schachter et al., 1987; Lansky et al., 1988), Germany (Reiss & Reiss, 1997),
Sweden (Levander & Schalling, 1988), South Africa (Le Roux, 1979), and
Turkey (Tan, 1988). The latter showed, however, higher percentage of right- and
left-handed women compared to men. More right-handedness in females and
more left-handedness in males were also reported in nonhuman animals, such
as cats (Tan, 1993b), chimpanzee (Corp & Byrne, 2004), and rats (Otmakhova,
 420 D. D. ELALMIS AND U. TAN

1989). Unfortunately, none of the authors studying the sex differences in

handedness considered the bodily measurements such as height and weight
of the participants.
Interestingly, the present study has found that the left-handers were heavier
and taller than the right-handers. This result indicates an association between
hand preference and body size. In accord, Tan (1992a) first reported significant
relations between body height and hand preference in right- and left-handers;
Mulligan et al. (2001) reported handedness differences in short and normal-
height subjects.
The dependence of sex difference in hand preference on body size was
proven in the present study by taking the individuals with nearly the same
heights and weights. In doing so, the sex difference in hand preference
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completely disappeared. This suggests a hormonal contribution to the origins

of hand preference: “hormones affecting body and brain size such as human
growth hormone and testosterone would also contribute to the development of
cerebral lateralization” (Tan, 1992a); “it may be that the hormones responsible
for growth and development also play some part in brain laterality. . .” (Mulligan
et al., 2001). Growth hormone was indeed shown to have significant relations
to grasp-reflex asymmetry in human neonates (Tan, 1994, 1995). The relation
of testosterone with handedness was also shown in human newborns (e.g., Tan
& Tan, 2001) and adults (e.g., Tan, 1991, 1992b, 1993a).

Hand Skill. In inter-subject analysis (mean PMTs for single individuals), sex
was not a significant factor for the RH-, LH-, and the LH–RH PMTs, in the first
and fifth trials of RH- and LH-writers. This is not consistent with Annett and
Kilshaw (1983), who found a sex difference in hand skill, but they measured
the peg-moving time by a stop watch instead of an algorhythm used in the
present work. Moreover, these authors took the mean of five trials; instead,
the present authors studied the single trials and single hand movements. Using
Grooved Pegboard Test, Schmidt et al. (2000) found that women were faster
than men. These authors did not consider any bodily measurements, but Peters
et al. (1990) showed that sex difference in hand skill (Purdue Pegboard) may be
confounded by sex differences in finger size: when thickness of index finger and
thumb were used as covariates, all significant sex differences in performance
completely disappeared. Therefore, considering also the present results, the
faster hands in women than men does not seem to be tenable.
The intra-subjects analysis (single hands) showed that sex was not a
significant factor for the RH- and LH-PMTs. So, there was not a female
advantage in hand speed, as in the inter-subject analysis. However, sex was
 HAND SKILL 421

found to be a significant factor for the LH–RH PMT, that is, difference
between hands in skill. Namely, the LH–RH PMT (faster right hand) was
significantly greater in women than in men for the first and fifth trials. This
was consistent with Annett and Kilshaw (1983), who found a right shift in the
LH–RH distribution of the PMTs in women compared to men. On the other
hand, this result also indicates that women are more lateralized than men in
asymmetric motor control of fine hand movements. This is consistent with the
notion that motor lateralization is more pronounced, in favor of the right hand,
in women than in men (e.g., Beric et al., 1997).
Some of the results considerably changed after taking the height as a
covariate. For instance, the mean LH-RH PMT was greater in women than in
men in the first trial of the RH-writers (inter-subject analysis), the difference
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being marginally significant (p = .06). This difference completely disappeared

under covariation of the height, however. Interestingly, the mean LH–RH
PMT was found to be equal (p = 1.00) in men and women after taking the
height as a covariate. In intra-subject analysis, in the first trial, sex was not a
significant factor for the mean RH- and LH-PMTs, but they were significant
if the height was chosen as a covariate, that is, the mean RH- and LH-PMTs
were significantly greater in men than in women. In the fifth trial, sex was not
significant for the RH-PMT, but nearly significant after taking the height as a
covariate, that is, the mean RH-PMT was greater in men than in women. Sex
was a significant factor for the LH-PMT, but it was not significant under height
as a covariate. These results indicate the importance of body size for analysis
of sex differences in handedness, in accord with Tan (1992a). The results of the
present work are not consistent with sex differences in hand skill; the previously
reported sex differences may depend on body size.

Right- and Left-Hand Skill in Right- and Left-Handers

Inter-Subject Analysis. Handedness for the first trial was found to be
a significant factor for the RH-PMT, which was significantly greater in
LH-writers than in RH-writers, but the mean LH-PMTs were not significantly
different in RH-writers and LH-writers. In the fifth trial, handedness was
significant for both of the RH- and LH-PMTs: the mean RH-PMT was
significantly greater in LH-writers than in RH-writers; the mean LH-PMT
was significantly greater in RH-writers than in LH-writers. In other words,
the right hand was faster in RH-writers than in LH-writers; the left hand was
faster in LH-writers than in RH-writers. Thus, the right-hand skill makes the
difference between the RH- and LH-writers, before practice. The expected
 422 D. D. ELALMIS AND U. TAN

difference between hand skills first emerged after practice, being faster right
hand and faster left hand in RH-writers and LH-writers, respectively. Neither
handedness group was superior to the other, being inconsistent with reports
about the faster hands in left-handers compared to right-handers (Kilshaw &
Annett, 1983).

Intra-Subject Analysis. In the first and fifth trials, handedness was significant
for the RH- and LH-PMTs: the right hand was faster in RH-writers than in
LH-writers; the left hand was faster in LH-writers than in RH-writers, as
expected. Neither group was superior. In inter-subject analysis, the superiority
of hand skills was different in the first and fifth trials, indicating the effects
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of practice on hand skill, depending on repeated trials. On the other hand,

considering the single hand movements yielded more conceivable results than
the intra-subject analysis.

Comparison of the First and Fifth Trials. There were significant differences
between hand skills in the unlearned first trials and the learned fifth trials. For
instance, in inter-and intra-subjects analysis, the mean RH- and LH-PMTs were
significantly greater in the first than the fifth trials. These results point out an
effect of practice on hand skill; the hands were faster in the fifth than the first
trials. This is in fact expected, because motor repetition causes motor learning.
This was recently shown in rats’ paw skill (Elalmis et al., 2003). Tan
(1989) previously reported that the mean PMTs linearly decreased at each
successive trials of the right and left hands. The present results are consistent
with this study. So, it may be concluded that repetition of hand movements
in the peg-moving task may cause motor learning exhibiting faster right- and
left-hands occurring at each successive trial (see Figure 6). It is indeed well
known that the repeated hand movements can gradually develop a stereotyped
pattern of agonist and antagonist muscle activity, inducing plastic changes in
the motor-related areas of brain, that is, motor learning (see Sanes, 2003).
Despite that, the mean LH–RH PMT did not show any significant changes in
RH- and LH-writers. So, the asymmetry in hand skill is not changed by practice.
However, in inter-subject analysis, the distribution of the LH–RH PMT in the
fifth trial shifted to the right compared to that in the first trial. The number of
individuals with faster right hand increased and the number of individuals with
faster left hand decreased in the fifth trial compared to those in the first trial. It
can be concluded from these results that the right- and left-hand skill improved
in favor of the right hand, depending on the motor leaning. So, analyzing the
 HAND SKILL 423

single trials in peg-placing task would be better than taking an overall average
for all the trials.

Effect of Repeated Hand Movements

It is expected that repeated hand movements in the peg-moving task may
induce a motor learning, which is well known to depend on practice (e.g.,
Willingham et al., 2001). Learning in peg-moving task in left-handers was
previously demonstrated by Tan (1989, 1993): the mean RH- and LH-PMTs
linearly decreased in the consequent five trials. In accord, Garry et al. (2003)
using Purdue pegboard found an improved performance with both hands over
the three trials with an increase in excitability of corticospinal motoneurones
controlling intrinsic hand muscles engaged in peg-placing task. Consistent with
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these results, the present work presents evidence for motor learning between
trials and even within a single trial. For instance, in RH-writers and LH-writers,
the mean RH- and LH-PMTs quadratically decreased with five consequent
trials, indicating faster right and left hands following practice. Moreover, the
percentage of individuals with faster left hand significantly decreased and the
percentage of individuals with faster right hand significantly increased in the
fifth trial compared to the first trial in RH-writers; the mean RH- and LH-PMTs
in the fifth trail were significantly less than those in the first trial, except the
LH-writers exhibiting the same pattern but with no statistical significance,
probably because of smaller sample size. The effect of practice could even
be seen within a single trial including nine hand movements: consequent RH-
and LH-PMTs significantly decreased from the first to the last hand movement
in RH-writers. The LH-writers did not exhibit similar motor learning. These
results suggest that the first trial as a relatively unlearnt peg-moving task would
be most suitable for analysis of hand skill.

Fluctuating Asymmetry in Hand Skill

In RH- and LH-writers, the LH–RH PMTs fluctuated between faster right
(LH–RH PMT > 0) and faster left hand (LH–RH PMT < 0) within a single trial.
The incidence of the mismatched negativity (LH–RH PMT < 0) in RH-writers
was 25.3%, which increased to 27.1% in the fifth trial, but the difference was not
significant. Similarly, in LH-writers, the numbers of the participants exhibiting
faster left hand in the first trial (41.4%) was not significantly different from the
numbers of the participants with faster left hand in the fifth trial (38.9%). Thus,
the amount of fluctuating asymmetry in hand skill did not change with fatigue
of the participants.
 424 D. D. ELALMIS AND U. TAN

Fluctuating asymmetry may also reflect fluctuations in attention. The

aforementioned result does not support this hypothesis. Moreover, the numbers
of mismatched negativities in RH-writers and mismatched positivities in
LH-writers were not significantly different within the first and last halves of a
single trial. This also does not support the earlier idea about attention deficits
in fluctuating asymmetry. If this was true, the amount of the mismatched
negativities in RH-writers and the amount of the mismatched positivities in
LH-writers would increase during the second halves of a single trial or during
the fifth trial compared to the first trial.
On the other hand, it is conceivable that the fluctuating asymmetries during
single hand movements may indeed reflect the normally occurring stochastic
fluctuations in cerebral attention system during single hand movements. In this
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context, one can experience sometimes dissociations from current goals and
immediate actions. These lapses in attention can occur in isolation, lasting only
a single trial, as West and Alain (2000) reported: “the neural system supporting
goal-directed action fluctuates in efficiency over time and these fluctuations
are an inherent property of this system” (West & Alain, 2000). So, the
fluctuating asymmetries in the asymmetric control of hand skill may be a normal
property of the brain, which may be more pronounced in certain disorders with
much more fluctuating attentions such as in dementia, Alzheimer disease, and
schizophrenia (Ballard et al., 2001). The fluctuating asymmetry in peg-moving
task may be used in clinical analysis of these disorders. Interestingly, Peters
(1994) has argued that “not only attention asymmetrically distributed to the two
hands but this is the essential prerequisite for skilled bimanual activity.” The
present study did not investigate any bimanual activity; the analysis restricted
only to single hand skills; the authors could not find any clues for contribution
of attention to asymmetric hand skill. The study found, however, that the
left-hand PMT showed significantly more fluctuations than the right hand in
men, women, RH-writers, and LH-writers. So, the fluctuating hand skill may
be caused by the right brain in right- and left-handers. These results suggest
attentional fluctuations in single hand movements, depending on the transient
changes in the right prefrontal cortex responsible for attentional processes (see
West & Alain, 2000). Despite the dependence of the fluctuations of single hand
movements on the right prefrontal cortex, the fluctuations in the LH–RH PMTs
were not dependent to either sex or handedness.
The number of the positive (LH-RH PMT> 0) and negative signs (LH-RH
PMT < 0) were estimated in RH-writers and LH-writers within the first
trial. It was found (see Figure 6) that the LH-writers exhibited the most
accentuated fluctuations and mismatched signs (72.7%) between 3 and 6
 HAND SKILL 425

positive LH-RH PMTs. This was significantly higher than the proportions
of the mismatched negativities (44.6%) in RH-writers. The proportion of the
individuals with the least mismatched signs and least fluctuations (positives
in LH-writers and negatives in RH-writers) was significantly higher in RH-
writers than in LH-writers. These results suggest that left-handers are most
vulnerable to fluctuating asymmetry with mismatched asymmetry in hand
skill. Further, the left-handers exhibited a prototype for fluctuating asymmetry,
because the LH–RH PMT was not significantly different from zero in these
individuals.
Whereas the LH-writers exhibited a prototype of fluctuating asymmetry
(RH minus LH PMT = zero), the RH-writers exhibited a directional asymmetry
in favor of a population level right-handedness in manual skill. There may be
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several factors playing a role in this shift to the right, such as genetics and
hormonal influences on the asymmetric brain development. With regard to
genetics, the fluctuating asymmetric distribution in LH-writers seems to be
associated with individuals having no right-shift factor (see Annett, 2002, p.
315). This is also compatible with the “chance” gene for fluctuating asymmetry
in McManus’ model (see McManus, 1985). The testosterone hypothesis
(Geschwind & Galaburda, 1985) does not predict the distribution of hand
skill. The evolution of asymmetric writing centers in the human brain may
also play a role in emergence of the directional asymmetry in humans in favor
of right-handedness, because the right-hand skill is directly associated with
writing. If so, the hand preference in non-human beings should not show a
directional asymmetry, but it is now known that there are population-level
right-handedness in non-human beings (e.g., Guven et al., 2003).

Genetic Influences on Handedness

Hand Preference. In children with right-handed mother and father (FS-),
90.7% were right-handed (RH-writer) and 9.3% were left-handed (LH-writer).
For a predominant right-handedness, right- and left-handed children should be
born from heterozygote right-handed mother and father (Rl × Rl) according
to Mendel’s laws. If the mother is left-handed and father right-handed, 50% of
children will be right-handed and another 50% left-handed. In this case, again to
Mendel’s laws, mother should be homozygote-recessive left-handed and father
should be heterozygote right-handed (ll × Rl). Thus, a random distribution
of hand preference (50:50) may also be determined genetically (McManus’
chance (C) gene, 1985; see also McManus & Bryden, 1992; Corballis, 1997).
In case of a right-handed mother and left-handed father, one has the same
 426 D. D. ELALMIS AND U. TAN

proportions of the right- and left-handers as in FS-individuals. This suggests

that right-handedness may be inherited from mother’s X-chromosomes.

Hand Skill. In FS-individuals, the mean LH-RH PMT was significantly greater
than zero, in favor of the right-hand skill. Similar to hand preference, there was
a chance distribution of the LH–RH PMTs in FS+1 (left-handed mother ×
right-handed father) individuals. This again suggests that a chance distribution
may be genetically determined. This result is consistent with McManus’
chance (C) gene (see MacManus, 1985), which can be localized in mother’s
X chromosomes. This expectation was supported by the results from FS+2
individuals (right-handed mother × left-handed father): the mean LH–RH PMT
was significantly greater than zero as in FS-individuals, but it was significantly
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greater in FS+2 than FS-individuals, in favor of the right-hand skill. This

suggests that right-handedness in mother may cause more right-handedness
in children. These results suggest that mother would play a major role in
determining the direction of asymmetric hand skill.
Concerning the right- and left-hand skill, there was no significant difference
between the RH-PMTs of the FS- and FS+1 individuals, whereas the mean
RH-PMT was significantly smaller in FS+1 than in FS+2 individuals. Thus, the
FS+2 individuals had faster right hand than FS- and FS+1 individuals; although
the father is left handed, the children use their right hand faster, following
right-handed mother. Concerning the left-hand skill, the FS+1 individuals had
faster left hand than FS- and FS+2 individuals. Thus, it may be concluded that
a left-handed mother is associated with a faster left hand in FS+1 individuals,
as is also proposed for the right-hand preference (see earlier). A left-handed
mother with right-handed father may also increase the chance of left-hand skill
with a chance distribution of the LH–RH skill. The results showed that a chance
distribution in right- minus left-hand skill may reflect left-hand preference in
writing. Sinistral mother was found to be linked mainly to the right-hand skill,
suggesting a left-brain control under genetic influences. The results suggest
a X-linked genetic inheritance of hand skill. In accord, Laval et al. (1998)
provided the first molecular evidence for linkage to relative hand skill (cerebral
asymmetry) on the X chromosome.

Stochastic Resonance Hypothesis of Cerebral Laterality

The results of the present study frequently pointed out the stochastic fluctuations
in hand-skill asymmetry even during a single trial. These may be due to
stochastic attentional fluctuations. There was, however, no sign for fatigue,
 HAND SKILL 427

because the degree of fluctuations did not change between trials, and between
the first and last halves of a trial. Left-handers exhibited a fluctuating asymmetry
with a zero mean in LH-RH PMTs. Such a real fluctuating asymmetry, that is,
“random deviation from symmetry on bilateral characters” (Palmer & Strobeck,
1986, p. 391), in hand skill is important for the developmental instability, which
may be induced by genetic factors (see Thoma et al., 2002). Accordingly this
study has shown a genetic origin of fluctuating asymmetry in hand skill, which
may be linked to a chance gene (see earlier).
The functional fluctuating asymmetry in asymmetric hand skill, that
is, fluctuations between RH–LH skill between faster right and left hands
during a single trial in peg-moving task, implies stochasting fuctuations
within the right and left cerebral hemispheres activities. That is, intermittent
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fluctuations in right- and left-hand skill with their asymmetric properties

suggest stochastic fluctuations within the brain, exhibiting stochastic-resonance
phenomena, whereby the response of a nonlinear system like brain to a weak
input signal can be optimized by the presence of a non-zero level of noise
(Wiesenfeld & Moss, 1995). The human brain can use externally added noise
for behavioral responses (behavioral stochastic resonance), at the cortical level
(Kitajo et al., 2003). Mechanisms of intrinsic stochastic resonance between
self-organized activities may be a general organizing principle for central
nervous system functions. Studies in recent years stressed the importance of
endogenously generated activity as a determining factor in the topology of
neural networks from early developmental stages and throughout life (Penn &
Shatz, 1999). The terms “endogenous,” “intrinsic,” and “spontaneous” imply
that the activity is generated from within the system, which are synonymous
to the term “self-organized.” The brain may be viewed as a self-organizing
nonlinear system. Resonance effects may be expected from the nonlinear
interconnectedness of neuronal networks within the brain. The prerequisite
for a system to exhibit stochastic resonance is a threshold, which should be
exceeded in order to activate the system. When the signal is not strong enough
to exceed the threshold, small amounts of noise added either to the system or the
signal may occasionally suffice to trigger activation. Therefore, this is called
“stochastic resonance” (Gammaitoni et al., 1998). Adding noise improved,
for instance, the ability of paddle fish to detect prey (Russell et al., 1999),
somtosensation (Collins et al., 1996), and the speed of memory retrieval (Usher
& Feingold, 2000). The endogenous activity of physiological systems may
play the role of “noise” for stochastic resonance-based processing of neuronal
signals (e.g., Balazsi et al., 2001; Glanz, 1997). The stochastic resonance
phenomena were studied separately within the right and left brain (Kitajo
 428 D. D. ELALMIS AND U. TAN

et al., 2003), but the interactions between two hemispheres. That is, noise into
the right hemisphere, signal into the left hamisphere, were not yet studied.
The results suggest stochastic resonance phenomena originating from the right
and left brain (stochastic fluctuations in the right- and left-hand skills), and
a possible interaction between right and left cerebral hemispheres (stochastic
fluctuations in the differences between the right- and left-hand skills). The
study has also demonstrated that the stochastic resonance phenomena may be
a genuine characteristic of the brain, which may have genetic origins.
It seems now to be well established that the brain exhibits stochastic
resonance phenomena within both the right and left cerebral hemispheres. This
principle can be applied to lateralization of action in brain. The stochastic
fluctuations in the right minus left hand skill apparently reflect the stochastic
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interactions between the stochastic resonance phenomena of the right and

left brains. A net asymmetry in the stochastic resonance phenomena of the
right and left cerebral hemispheres may result from a genetically determined
asymmetric coupling between hemispheres, in favor of the left brain in
right-handers. The bi-hemispheric coupling is still stochastic, but the net
effect will be in favor of the left brain because of genetically determined
properties of neuronal networks. The direction of stochastic coupling between
the stochastic resonances of both cerebral hemispheres will depend on the
predominant net effect of one of the interacting oscillators, according to
coupled oscillators theory (Abarbanel, 1996). A purely stochastic coupling
between cerebral hemispheres will establish stochastic bi-directional resonance
in bi-hemispheric individuals. The former may create a predominantly right-
handed persons, whereas the latter may create left-handed persons with no net
resonance effect between hemispheres. As mentioned earlier, both situations
may depend on the genetically determined chaotic properties of the right and
left cerebral hemispheres.

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