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The Hoffmann Reflex from the Flexor Pollicis Longus of the thumb in Left-
Handed Subjects: Spinal Motor Asymmetry and Supraspinal Facilitation to
Cattell's Intelligence Test
Üner Tan a
a
Atatürk University, Medical Faculty, Institute of Physiology, Erzurum, Turkey

Online Publication Date: 01 October 1989

To cite this Article Tan, Üner(1989)'The Hoffmann Reflex from the Flexor Pollicis Longus of the thumb in Left-Handed Subjects: Spinal
Motor Asymmetry and Supraspinal Facilitation to Cattell's Intelligence Test',International Journal of Neuroscience,48:3,255 — 269
To link to this Article: DOI: 10.3109/00207458909002169
URL: http://dx.doi.org/10.3109/00207458909002169

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 Intern. J . Neuroscience, 1989. Vol. 48, pp. 255-269 1989 Gordon and Breach, Science Publishers, Inc.
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THE HOFFMANN REFLEX FROM THE FLEXOR

POLLICIS LONGUS OF THE THUMB IN
LEFT-HANDED SUBJECTS: SPINAL MOTOR
ASYMMETRY AND SUPRASPINAL FACILITATION
TO CATTELL’S INTELLIGENCE TEST
UNER TAN
Atatiirk Univer.yity, Medical Faculty, Institute of Physiology, Erzurum, Turkey

i Received Muq’ 3 I , 1989)

The spinal motor asymmt:try was studied in left-handers. Hand preference was assessed by Geschwind
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scores (GS), and hand skill by peg moving task. the reflex responses were recorded from the long flexor
muscle of the thumb. The H-reflex could be elicited by averaging during cortico-spinal facilitation (volun-
tary insometric force). The Cattell’s Culture Fair Intelligence Test was used to assess the individual
differences in mental abilities. H-reflexes were found to be significantly larger on the left than the right side.
The amplitude of H-reflex increased linearly with force applied to transducer by the thumb. This facilitation
was more pronounced for the left than the right reflexes. Removal of the visual input caused facilitation
in H-reflex (supraspnal disinhibition). Post-activation potentiation was also observed in H-reflex. There
was a positive linear correlation between the degree of left-hand preference (-GSs) and left-hand skill. The
correlation for the right-hand skill was not significant. There was a linear correlation between the degree
of left-hand preference and the right minus left hand skill. There was an inverse correlation between
left-hand skill and H-reflex from left. The correlation for the right side did not reach the 5% significance
level. The force-reflex relarion did not show any significant change to IQ for the right H-reflex. The left
H-reflexes were significantly larger in subjects with high IQ than those with low IQ. The regression line and
its slope for the force-reflex relation on the left was found to be higher in subjects with high IQ than those
with low IQs. There was a positive linear relationship between IQ and H reflex from left. The correlation
for the right side was not ds pronounced as that for the left side. The left minus right H reflex was also
positive linearly correlated with IQ. These results provide further evidence for the psycho-motor hypothesis
(Tan, 1988b) as well as the spinal motor asymmetry to handedness. It was suggested that lateralization of
cognitive and motor functions would be essential to create subjects with high psychomotor capacity.

Keywords: H-reflex, licmde~lnr.v.\,Iuterulkaiion, motor neuron, spinal cord, intelligence

INTRODUCTION

I have previously provided evidence for a spinal motor lateralization in cats (Tan,
1984). This asymmetry was observed in both the intact and spinal animals. The right
and left dominance in motoneuronal excitability assessed by the H-reflex recovery
curve from the gastrocnemius nerves did not change after spinalization. However, a
right or left dominance appeared in ambilateral cats following spinalization. There
was no correlation between paw preference and spinal motor asymmetry. According-
ly, Goode et al. (1980) have reported that seven subjects had higher H-reflex recovery
curves in the right leg, and four in the left leg, although all of these subjects expressed
right hand preferencc.
Tan (1985a, b) has later shown that there is indeed an inverse relationship between
handedness and the excitability of motoneurons innervating the postural soleus
muscle in right-, and left-handed subjects, i.e., the excitability of these motoneurons
was higher on the left than the right side in right handers, and vice versa in left

255
 256 U TAN

handers. To explain these unexpected results, I have suggested that the left leg takes
over the support function (postural adjustments) so that the right foot may gain
freedom to perform skilled movements in right-footed persons. Presumably. the
inhibitory action of the pyramidal system onto the antigravity muscles is more
pronounced on the preferred side (operative side) than the nonpreferred side (suppor-
tive side). Such an organization would be essential for fine motor control, since
inhibition is essential for fine regulation of the motor acts. Accordingly, I have
recently found that the H-reflex amplitude is inversely correlated with hand skill. and
suggested that small reflexes are especially suitable for the fine motor control of the
rapid aimed-movements (Tan, in press a, b).
With regard to hand skill, it is to be expected that the excitability of motoneurons
innervating the flexor hand muscles should be higher on the preferred side than the
nonpreffered side. In accord with this proposal. 1 have found that the excitability of
motoneurons-the size and recovery curve of the H-reflex-innervating the wrist
flexors is higher on the preferred side than the nonpreferred side in right-. and
left-handed subjects (Tan, in press a,b). In the present work, the above hypothesis was
further tested on the long flexor muscle of the thumb in left-handed subjects, since the
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thumb-finger grip is of importance in writing. which shows the most significant

association with hand preference. Moreover, the functional value of the thumb is
about half that of the whole hand. The size of the H-reflex was used to assess the
spinal motor dominance in left-handed subjects. The properties of the H-reflex were
studied in relation to supraspinal facilitation and intelligence in these subjects. A
preliminary account of these experiments has been presented elsewhere (Tan, in press
C).

METHODS

Suhjec ts
The experiments were performed on twenty left-handed subjects of both sexes drawn
from the students of the Medical Faculty. Their ages ranged from 18 to 23. None
declined to participate in the study. They were healthy, devoid of neurological or
psychiatric sigr,s, and symptoms.

Hand Prcftvcnce and Hand Skill

Hand pwference. The hand preference was assessed by the Edinburgh Handedness
Inventory (Oldfield, I97 1). The left-hand writing was considered essential for left-
handedness. Geschwind scores (see Tan. 1988a) smaller than zero (-GSs) indicated
left-handedness (non-right handedness); there were various degrees of the left hand
preference according to different GSs.

Hand skill. To evaluate the hand skill, the Purdue Pegboard (Purdue Research
Foundation, 1948) was used, which was modified for accurate time measurements and
a more difficult task. The peg board consisted of two parallel rows of 25 holes. There
were 25 loose-fitting dowelling pegs in one row. The subjects were required to shift
these pegs to the corresponding holes as fast as possible. Pegs were moved from right
to left with the right hand, and then from left to right using the left hand. An electronic
device measured the exact time elapsed between the first and last peg movings for each
hand separately. One trial consisted of the time elapsed to move 25 pegs with one
 H-R EFLEX A N D INTELLIGENCE IN LEFT-HANDERS 257

hand. Ten trials were performed by each hand. The mean peg moving times were
calculated for each hand. The significance for the difference between the mean peg
moving times of both hands was tested by the Student’s t-test for paired data. Some
left-handers (left-hand writing and GSs < 0) proved to be ambidexters in peg moving
task (hand skill).

Intelligence
Individual differences in mental ability were established by using the Cattell’s Culture
Faire Intelligence Test (see Tan, 1988b). In the present work, scale 2 was used as a
group test. The row scores were converted into IQs. Scale 2 consists of the following
for tests: 1 . Series: Select the item that complete the series. 2. Classification: Mark the
one item in each row that does not belong with the others. 3. Matrices: Mark the item
that correctly completes the given matrix or pattern. 4.Conditions: Insert a dot in one
of the alternative design.
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Hofmann Reflex
The recordings were made from the long flexor muscle of the thumb of both hands.
The subjects were sitting comforably with their hands on the table. The fingers of the
hand studied were wrapped loosely. The active (recording) electrode was placed over
the belly of the flexor pollicis longus muscle and the reference electrode over the
proximal phalanx of the thumb. The electrical stimulation was performed by stimulat-
ing electrodes placed and wrapped over the median nerve at the wrist. The ground
electrode was a sheet metal taped to the arm just above the elbow. The cathode was
proximal to anode. The stimuli (0.2 ms pulses) were delivered by a constant current
stimulator at the frequency of I/s. The stimulus intensity was just at threshold for
motor fibres of the median nerve, so that it was always tried to create the same
stimulus conditions for both hands.
The H-reflex could only be elicited by facilitation of the motoneurons through the
voluntary impulses onto the motoneurons. To elicit the H-reflex, the subject was
instructed to press the ball of the thumb against a flat circular brass disk attached to
an isometric force transducer. The force exerted by the subject’s thumb was made
visible on a digital voltmeter connected to the force transducer, so that the subject
could control and maintain the required force by visual feedback. The threshold force
for the H-reflex was kept constant for all subjects and for both hands.
At the threshold force, the median nerve was stimulated by single shocks at an
intensity just threshold for motor fibres. To make the H-reflex visible, EMG signals
were averaged over 128 trials, while the subject exerted a constant force on the strain
gauge. Following a resting period of one minute, the subject was asked to double the
force, and the same procedure as before was repeated under the same stimulus
conditions. The responses were also recorded at three and four times threshold forces.

TABLE 1
The mean reflex latencies (ms) from the long flexor muscle of the right and left thumbs

Muscle H-reflex S.D. LL(1) S.D. LL(2) S.D.

Left FPL 29.9 2.8 48.8 5.7 68.6 4.6
Right FPL 30.4 2.6 52.3 8.6 71.2 8.8

FPL: flexor pollicis longus; LL(1): the first long latency reflex; LL(2) the second long latency reflex.
 258 U. TAN

TABLE 2
The mean H-reflex amplitudes (mV) from the long flexor muscles of the right and left thumbs

H-reflex Mean S.D. minimum maximum median

Right 0.16 0.08 0.04 0.44 0.16

Left 0.36 0.16 0.04 0.82 0.33
L-R 0.20 0.18 - 0.04 0.75 0.15

L-R: left minus right reflex amplitude

The latencies of the reflex responses were measured by the cursor of the oscilloscope.
The responses were plotted on the paper by a plotter connected to the oscilloscope.
The peak-to-peak amplitudes of the H-reflexes were measured by callipers on the
paper; the metric values were converted to voltages.
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A.LEFT B.RIGHT

3 d u &'
- - ' r- ,---./- -3

FIGURE 1 Reflex responses from the long flexor muscle of the right and left thumbs in a left-handed
subject. A1,2 and B1,2 refer to the reflex responses and reflex contractions from the left and right sides,
respectively. The force applied to the force transducer was one times threshold (a certain force eliciting an
H-reflex in all subjects). S , stimulus artefact; M, M response; H, H-reflex. In A3,4 and B3,4 the force was
two times threshold. In AS,6 and B5,6 the force was three times threshold.
 H-REFLEX AND INTELLIGENCE IN LEFT-HANDERS 259
RESULTS

Reflex Responses from the Long Flexor Muscles of the Right and left Thumbs
Table 1 shows the mean latencies of the reflex responses from the right and left flexor
pollicis longus muscles. The reflex latencies for the right side seems to be longer than
those from the left side.
Table 2 shows the mean amplitudes of the H-reflexes from the long flexor muscles
of the right and left thumbs. The mean amplitude of the H-reflex from the left side
was significantly larger than that form the right side ( t = 3.90, df = 19, p = .001).
The original records were presented in Figure 1. The records in A and B are due
to the reflex responses from the long flexor muscles of the left and right thumbs,
respectively. This subject (GS = -60) was ambidextrous in peg moving task. His
father, mother, and grandfathers were also left-handed. The latencies of the responses
in B1 (the first and subsequent arrows) were 26, 35, 42, 46, 50, 62, and 69ms. The
arrows in B4 show the latencies of the reflex contraction of the muscle, which began
at 40 ms (first arrow), had a peak at 53 ms (second arrow), and ended at 64 ms (third
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arrow) following stimulation of the median nerve. The latencies in A5 (the first and
subsequent arrows) were 27, 31, 35, 42, 47, and 72ms. the reflex contraction of the
muscle began at 34 ms following stimulation of the median nerve.
The reflex responses in A and B were recorded at 1, 2, and 3 times threshold force
applications. It is clearly seen that the responses, especially the H-reflexes, were larger
on the left than the right side, and so were the reflex contractions of the muscle.
Figure 2 presents the relationship between the force applied to the isometric force
transducer and the amplitude of the H-reflex from the right (dashed line) and left
(straight line) sides. The relations were strictly linear (r = .99). The slope for the
straight line (left reflex) was higher than that for the dashed line (right side). More-
over, the mean reflex amplitudes for the left side were larger than those for the right

0.4 -
(mV)

0.3

'
0.2

0.1

0.0 -
1 2 3 (xT) 4
FIGURE 2 The relationship between force and the H-reflex from the long flexor muscle of the right and
left thumbs in the total sample (N = 20). Abscissa: force applied to the force transducer as times threshold
(xT).Ordinate: mean amplitude of the H-reflex (mv). The standard deviations were omitted because of
+
simplicity. Dashed line: y = 0.006 O.O4x, r = .99. Straight line: y = 0.04 + O.O8x, r = .99).
 260 U TAN

side. The differences between the mean right and left reflexes for each force level
ranging from 1 x to 4 x threshold were found to be statistically significant at
p = .005 level.
Figure 3 presents another example for the left-dominant H-reflex in left-handers.
The H-reflex (H) in A and B were recorded from the long flexor muscles of the left
and right thumbs, respectively. It is seen that the H-reflexes at increased force levels
(records I , 2, and 3) are much larger on the left than the right side. This Figure was
presented to show the supraspinal influence onto the motoneurons. In records 4.the
subject was required to close the eyes while he applied the same force as in records
3 (3 x threshold force). This procedure caused a prominent facilitation in the reflex
responses of both sides (A4 and B4). The first and following arrows indicate the
latencies of the responses. In A4, these latencies were 27, 30, 34, 45, 50, 63, 120. and
134 ms; in B4, 26, 30, 33, 35, 44, 50, 58, and 68 ms.
The records in Figure 4 illustrate the post-activation potentiation in the H-reflex.
The records 1 in A, B, C, and D were taken at 1, 2 , 3, and 4 times threshold forces.
The H-reflexes increased in size as usual, at each increment of force. Without resting
for one minute following this first trial, the whole procedure was repeated (records 2
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in A, B, C, and D). It is clearly seen that the H-reflexes were much larger now,
indicating a post-activation potentiation.

Relationship Between Hand Preference and Hand Skill

The hand preference was assessed by GSs, and hand skill by the peg moving task. The
diagram above in Figure 5 shows the relationship between GS and the stabilized peg
moving time (last trial) for the right (open circles, dashed line) and left hand (closed
circles, straight line). There was a negative linear correlation between GS and the peg
moving time, i.e., the degree of the left-hand preference (-GS) increased as the peg
moving time decreased. In other words, the hand skill increased as the degree of
left-handedness increased, indicating a postive linear correlation between hand skill
and left-hand preference, but the correlation was significant only for the left hand

A.LEFT 8.R I G H T
1 1
M H

A
.+ 3 - 3
I

\ 4 d"i"
1 - L

I r/""* A

5 0 ms

FIGURE 3 Supraspinal facilitation of the H-reflex by removing the visual input. M, H: M and H
responses from the long flexor muscle of the left (A) and right (B) thumbs. Records I , 2, and 3 are taken
at 1, 2, and 3 times threshold force applications, respectively. Records 4 in A and B: 3xT, as in record 3,
but the eyes were closed.
 H-REFLEX AND INTELLIGENCE IN LEFT-HANDERS 26 1

-a
h-
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1
,
60ms

FIGURE 4 Post-activation potentiation in the H-reflex. M, H: M and H responses. A, B, C, and D:

responses from the long flexor muscle of the right thumb, at Ix, 2x, 3x, and 4xT force applications,
respectively. Records 1. first trial; records 2, second trial. There was not a resting period of one minute
between the first and second trials. Latencies: start of the H-reflex = 23ms; L1 = 43ms; L2 = 60ms;
L3 = 105ms.

(r = .49,p < .05). The dotted line is the regression line fitted to the mean peg moving
times (10 trials) for the right hand. The correlation was not significant ( r = .28,
p > .05). There was a negative linear correlation between the mean peg moving time
for the left hand and GS. This correlation was found to be statistically significant
(r = .58,p < .02).
The diagram below in Figure 5 shows the relationship between the right minus left
peg moving time and the degree of left-handedness (-GSs). There was a positive linear
correlation between the left minus right values of the mean peg moving times and GSs
(open circles, dashed line). The correlation was found to be statistically significant
(r = .60, p < .05). The same significant correlation was found when stabilized (last
trials) peg moving times were considered (closed circles, straight line).

Relationship Between Hand Skill and H Reflex

Figure 6 shows the relationship between the mean H-reflex (mV) and peg moving
time. There was a positive linear correlation between the mean peg moving times for
the right hand and the H-reflex from the right side (open circles, dashed line), but the
correlation did not reach the 5% significance level (r = .44, p = .lo). There was a
 262

36 -
0
(S)

-
*.....-*. ........
0
32
0 0
................... 8*,./'"
............... - - -/-&= - --
0

0 .a-..... - -
28-
; ._--- -.-
-y.-c .
,.i-6-c
./a

24 -
/. #.C./

0 0
0 0

e .

e e
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20 1

0 8

-100 - 80 - 60 -4 0 -20 (GS) 0

FIGURE 5 The relationship between the hand preference ( G S ) and hand skill (peg moving). Absassa:
left-hand preference (-GS); ordinate: peg moving time (s). Above: open circles, dashed line refer to the
stabilized peg moving time (last trial) for the right hand versus GS (y = 29.7 + 0 . 0 3 ~r. = .22, p > .05);
closed circles, straight line: the stabilized peg moving time for the left hand versus GS (y = 28.1 + 0 . 0 5 ~ .
r = .49, p < .05): dotted line: the mean peg moving time (10 trials) for the right hand versus GS
(y = 3 1.8 + 0 . 0 4 ~r. = .28. p > .05);interrupted line: the mean peg moving time for the left hand Lersus
GS (y = 30.0 + 0 . 0 6 ~r. = .5X. p < .02).

positive linear correlation between the mean peg moving time for the left hand and
the H-reflex amplitude from the left side (closed circles, straight line). The correlation
was found to be statistically significant (1. = .59. p < .0.02). The slope for the straight
line was much higher than that for the dashed line.

Relationship Between H R e f e x and IQ

The mean IQ was found to be 124 in the total sample ( N = 20). Therefore. the
 H-REFLEX A N D INTELLIGENCE IN LEFT-HANDERS 263

-.-22 24 26 28 30 32 34
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PEG MOVING TIME ( s )

FIGURE 6 The relationship between hand skill and H-reflex for the right and left hands. Abscissa: the
mean peg moving time (s 1; ordinate: H-reflex (mV). Dashed line, open circles; peg moving time for the right
hand versus H-reflex from the right side (y = -0.09 + O.O06x, r = .44, p = .lo). Straight line, closed
circles: peg moving time lbr the left hand versus H-reflex from the left side (y = - 0.8 + O.O4x, r = .59,
p < .02).

subjects were divided into two subgroups as the subjects with low (below 124) and
high IQ (124 and above). Figure 7 shows the relationship between the force applied
to the transducer by the thumb and the size of the H-reflex to force in subjects with
low and high IQs. For the subjects with low IQs, the H-reflex for the right side was

h
0.6 -
>
E
L d

0.2 '

0.0

FIGURE 7 The relationship between H-reflex and force in subjects with low and high IQ. Abscissa:
threshold force applied to the force transducer by the thumb. Ordinate: H-reflex amplitude (mv). Dashed
line, open circles: H-reflex from the right side versus force in subjects with low IQ (y = 0.004 + O.O3x,
r = .99). Straight line, closed circles: H-reflex from the right side versus IQ in subjects with high IQ
(y = - 0.01 + 0 . 0 5 ~ .r = .99). Dotted line, open triangles: H-reflex from the left side versus IQ in
subjects with low IQ (y = - 0.006 +
O.O6x, r = .99). Straight line, closed traingles: H-reflex from the left
side versus IQ in subjects with high IQ (y =0.05 + O.Ix, r = 1.0).
 264 U. TAN

correlated with the force well (open circles, dashed line, r = .99). In the subjects with
high IQs, the H-reflex from the right side versus force correlation was found to be
exact (closed circles, straight line); the reflex amplitudes were slightly larger than those
for the formers.
For the H-reflexes from the left side, there was an exact correlation between the
H-reflex amplitude and force in the subjects with low IQs (open triangles, dotted line).
In the subjects with high IQs, there was a positive linear correlation between the force
and H-reflex amplitude (Y = 1.0). The slope of this regression line was much higher
than that for the left H-reflex of the subjects with low IQs. Moreover, the mean reflex
amplitudes from the left side was larger in subjects with high IQs (closed traingles) than
those for the subjects with low IQs (open triangles). The higher slope of the regression
line was also found for the subjects with high IQs and right-hand reflexes. but the
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-,--A A'..' 8A E o o
0.0 .A

9(3 100 110 120 130 140 150 7-

160
IW

w 0.4

' I

= o'2
0.0
90
L 100 110 120 130 140 150
IQ
160
I

FIGURE 8 The relationship between H-reflex and IQ. Above diagram: abscissa, IQ; ordinate, maximum
H-reflex (mV). Dashed line, open circles: H-reflex from the right side versus IQ (y = - 0.4 + 0.004~.
r = .50,p < .05). Straight line, closed circles: H-reflex from the left side versus IQ (y = - 0.8 + O.OIx,
r = .58, p < .01). Dotted line, open triangles: left minus right H-reflex versus IQ (y = - 1.0 + O.OIx,
r = .68, p < .01). Below: abscissa, IQ; ordinate, the mean H-reflex for all force levels. Dashed line. open
circles: H-reflex from the right side versus IQ (y = - 0.1 + 0.002x, r = .38, p > .05). Straight line,
closed circles: H-reflex from the left side versus IQ (y = - 0.7 + O.O07x, r = .65, p < .01). Dotted line,
open triangles: left minus right H-Reflex versus IQ (y = - 0.6 + O.O07x, r = .62, p < .01).
 H-REFLEX AND INTELLIGENCE IN LEFT-HANDERS 265

difference between the slopes of the regression lines for the left reflex of the subjects
with the low and high IQs were much more pronounced than the right reflex.
The diagram above in Figure 8 shows the relationship between the maximal
H-reflex and IQ. There was a positive linear correlation between the maximum
H-reflex from the right side and IQ (open circles, dashed line). The correlation was
found to be significant ( r = S 0 , p < .05). There was also a positive linear correlation
between the maximum H-reflex from the left side and IQ (closed circles, straight line).
The correlation was found to be statistically significant at a higher significance level
than for the right side ( r = .58, p < .Ol). The slope for the straight line was found
to be higher than the dashed line. The left minus right H-reflex amplitude was also
positive linearly correlated with IQ (open triangles, dotted line). The slopes of the
straight line and dotted line were the same.
In the diagram below in Figure 8, the mean reflex amplitudes from I , 2, 3, and 4
times threshold force applications were taken instead of the maximal amplitude of the
H-reflex. There was a positive linear correlation between the mean H-reflex from the
right side and IQ (open circles, dashed line), but the correlation did not reach the 5%
significance level ( r = .38, p < .05). The mean H-reflex from the left side was also
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positive linearly correlated with IQ (closed circles, straight line). The correlation was
found to be highly significant ( I = .65, p < .01). The slope for the straight line was
much higher than that for the dashed line. The left minus right H-reflex was also
correlated with IQ well (open triangles, dotted line, r = .62, p < .Ol).

DISCUSSION

Refrex Responses from Long Flexor Muscles of the Left and Right Thumbs
Under isometric conditions, essentially three responses were recorded in most of the
subjects. The first one was the Hoffmann (H) reflex. The overall reflex latencies
seemed to be shorter for the left than the right side, but the differences were not found
to be statistically significant. These latencies were very close to those frequently
reported in the literature. Interestingly, the mean delay between the H-reflex and the
first long-latency reflex was found to be 18.9ms, and between the first and second
long-latency responses 19.8 ms for the left side. For the right side, they were 21.9, and
18.9 ms, respectively. Spinal and/or transcortical pathways have been suggested to
induce these long-latency responses. Deuschl et al. (1985) have assumed a central
delay of at least 18.8 nis for a transcortical route. Therefore, the long-latency respon-
ses would fit with a transcortical pathway. The slightly shorter latencies for the left
side could be one of the features of faster left hand in left-handers. The mean
H-reflexes were found to be significantly larger on the left than the right side. Thus,
the motoneuronal excitability was higher on the left than the right side. These strong
reflexes from the left flexor pollicis longus muscle would be essential for the strong
thumb-finger grips.
Interestingly, there was an exact positive linear correlation between the H-reflex
amplitude and the force level. This exact correlation in the force-reflex relation is an
example for the integrative function of the nervous system, i.e., algebraic summation
at the motoneuronal membrane, since the same number of motoneurons were re-
cruited at each force level. The subject exerted an isometric force to the transducer
voluntarily. This induced a motoneuronal facilitation originating from supraspinal
structures and also operating within the gamma motoneuronal spindle loop. The
cooperative facilitatory effects of these corticomotoneuronal and gamma-spindle
 266 U . TAN

systems created a recruitment of motoneurons so that equal numbers of motoneurons

were added at each step of force. The force-reflex relation was more pronounced on
the left than the right side, indicating that, at equal increments of the force level, more
motoneurons were recruited on the left than the right side. In other words, the
motoneurons on the left (preferred hand) are more susceptible to facilitation than
those on the right (nonpreferred hand). This higher motoneuronal excitability could
be the origin of the left-hand preference as a driving force for left-handers to use the
left hand preferentially.
Removal of the visual input by closing the eyes caused a considerable increase in
H-reflex. Thus, the supraspinal input originating from visual cortex is in fact inhibit-
ory for the motoneurons studied. Apparently, closing the eyes removed this inhibition
and caused an increase in the number of motoneurons fired. It is well known that the
aimed fine movements are better executed under visual guidance (visuomotor con-
trol). It is, however, conceivable that fine motor control needs fine inhibitory regula-
tion. Therefore, the apparently inhibitory effect of the visual system on the mo-
toneuronal activity seems, in fact, to be advantagous for fine motor control. Accor-
dingly, I have recently found that hand skill is inversely correlated with reflex size
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(Tan, in press b).

Hand Preference and Hand Skill

The degree of hand preference (-GSs) was correlated only with left hand skill, which
increased with left hand preference linearly. Thus, GSs can be taken as index for left
hand skill in left handers. Annett (1970) has reported that asymmetries in hand
preferences and skill are related. She clasified the school children in to groups of pure
left-, mixed left-, mixed right-, and pure right-handers. The mean differences between
hands on the peg moving task were plotted for these groups; the relationship was
found to be positive linear, i.e., the skill difference between hands was most pronoun-
ced in subjects with pure left-, and pure right-handedness. However, she did not show
the leading role of the left hand, for instance, in determining the degree of left hand
preference. In the present work, it was also shown that there was a positive linear
correlation between the right minus left hand peg moving time and the degree of left
hand preference. The difference between hand skills on the peg moving task increased
as the left hand preference (-GSs) increased linearly.

Hand Skill and H-Reflex

Similar to correlation between left hand skill and left hand preference, only the left
hand skill was correlated with H-reflex from the left side. The mean peg moving time
for the left hand increased linearly with the amplitude of the H-reflex from the left
side. That is, there was a negative linear correlation between left hand skill and
H-reflex, i.e., higher skill was associated with small reflex. The same correlation was
also found for the H-reflex from the wrist flexors (Tan, in press a, b). As mentioned
in these recent works, a large reflex is not associated with fine motor control such as
the peg moving task. For instance, the fine visuomotor control is probably processed
by predominantly inhibitory inputs originating from the visual system, as discussed
above. it is well known that small motor units (small reflex outputs) are associated
with fine motor control, and the large ones (large reflex outputs) with gross move-
ments. Therefore, it is not surprising that small reflexes were found to be associated
with higher hand skill, although this was only true for the left hand. This indicates that
the left hand of left-handers is under a more precise central control than the right
 H-REFLEX AND INTELLIGENCE IN LEFT-HANDERS 267

hand. The higher slope of the straight line for the left hand skill-reflex relation
supports this conclusion.

H-Reflex and ZQ
A close relationship was found between IQ and H-reflex. This was again more
pronounced for the left than the right hand. The mean reflex amplitudes from the left
side were found to be significantly larger in subjects with high IQ than those with low
IQ. The slope for the IQ-reflex relation was found to be higher in subjects with high
IQ than those with low IQ. This result is accordance with psychomotor hypothesis
(Tan, 1988b), suggesting a strong interrelationship between cognitive and motor
functions.
The slope of the left force-reflex line was higher in subjects with high IQ than those
with low IQ. Thus, the supraspinal reflex facilitation was more pronounced in the
subjects with high IQ than those with low IQ. High IQ was associated with large
reflexes (high motoneuronal excitability) on the preferred side. I have recently shown
in left handers that motor learning on the peg moving task is better in subjects with
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high IQ than those with low IQ, but this was true only for the right hand (Tan, 1989b).
Moreover, the right hand skill was found better in subjects with high IQ than those
with low IQ. It was concluded from these results that the left cerebral hemisphere
would be of importance for the fluid intelligence (Cattell, 1971). The significant
association of the H-reflex with IQ established in this work suggest that the right
hemisphere might contribute to the cognitive output of the brain in left-handers. In
fact, it is to be expected that the preferred hand should be easily accessible for quick
motor actions to perform cognitive motor tasks such as writing. It can also be argued
that the right hand could have a higher plasticity in motor learning than the left hand
in left-handers (Tan, 1989b). This is probably because the left hand has already
acquired a high motor skill by exercise and motor dominance of the left brain.
However, there was no significant difference between the learning curves of the right
and left hands in these subjects (Tan, 1989b).
The previous results obtained from manual proficiency (Tan, 1989a) and hand skill
(Tan, 1989b) suggested a prominent role of the left hemisphere in the fluid intelligence.
The results of the present work showed now that the right hemisphere could also be
of importance for cognitive output of the brain, provided that the spinal cord does
not possess its own cognitive style. In previous works, the left hemisphere superiority
in cognition was associated with its analytical and language capacities. The right
hemisphere could contribute to cognition by triggering other functions which would
enhance the cognitive efficiency of the left hemisphere.
The more pronounced reflex asymmetry observed in bright subjects suggests a more
lateralized brain and spinal cord in these subjects. The IQ-reflex relation indicates in
fact that intelligence is not in possession of the brain alone; the spinal cord is also in
close association with cognitive functions. Sheehan and Smith (1986) reported that
increased lateralization of cerebral organization is associated with superior perfor-
mance on measures of spatial reasoning. Rhodes et al. (1969) found that bright
children have larger visual evoked potentials than dull children; dull children did not
demonstrate hemispheric asymmetry, and the evoked responses of the bright children
are consistently of larger amplitude over the right than over the left hemisphere.
Richlin et al. (1973) have also found that there is no asymmetry in auditory evoked
cortical response in the retarded sinistrals.
Rhodes et al. (1969) suggested that the larger evoked responses in bright children
would be associated with the level of attention. According to Lindsley (1957) the dull
 268 U TAN

child’s lower responsiveness, compared to normal and bright subjects, indicates a

disfunctioning reticular activating system. Accordingly, it is possible that reticular
activating system functioning well in subjects with high IQ could exert a more
pronounced facilitatory effect on the spinal reflex response than that in subjects with
low IQ. In attention demanding tasks, the cerebral blood flow was found to be greater
in the right frontal regions than the left ones (Deutsch et al., 1987). Jung et a]. (1984)
have clearly shown by recording slow brain potentials that the left hemisphere is
dominant for language and calculation in the vast majority (86.7%) of left handers.
The recent works (Tan, 1989a, b) indicated that the left hemisphere is of utmost
importance in the Cattell’s intelligence test in left handers. Then the larger reflex
excitability on the left hand or left handers with high IQs may be due to the very
general role of the right hemisphere in attention and imagination. Then a cooperative
functioning of the brain hemispheres seems to be essential for an efficient cognitive
output of the brain. Giannitrapani (1985) has studied the frequency components of
steady state EEG and concluded that the overall activity is overwelmingly homolo-
gously symmetrical in the execution of higher cortical functions. However, he did not
analyzed the motor components of the cognitive functions. Therefore, he might have
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missed the actually existing brain asymmetry in cognition. The lateralized motor
asymmetries in cognition revealed by using different approaches (hand skill, manual
proficiency, spinal reflex) suggest that both hemispheres may contribute to cognitive
output of the brain including the spinal cord, but by using different mechanisms.
There was a positive linear relationship between the H-reflex size and IQ. This
correlation was much more pronounced for the left side than the right side. Here
again, we see the importance of the motor output in relation to cognition. The reflex
excitability is larger and the IQ-reflex relation is more pronounced on the preferred
side than the nonpreferred side in conjunction with high IQ, since the cognitive output
of the brain is expressed by the left hand in left handers. It can thus be concluded that
the cognitive processing takes place in the left hemisphere, the cognitive output of this
hemisphere is transmitted to the motor-dominant right hemisphere, transformed to
motor commands for the motor expression, which is then executed by the left hand
possessing higher motor skill than the right hand in left handers. It can be suggested
that the cognitive and motor processings do not occur within the same hemisphere.
Such an organization would be most advantageous for economy, precision, and
efficiency of the brain in execution of cognitive functions in conjunction with motor
performance. This lateralization of cognitive and motor functions would be essential
to create subjects with high intellectual capacity. Indeed, it was also found that the
left minus right reflex size was positive linearly related to intelligence.

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