JOURNALOFNEUROPHYSIOLOGY
Vol. 44, No. 1, July 1980. Printed in U.S.A.
Mechanical and Neural Feedback Factors in
Postural Hand Tremor of Normal Subjects
ROBERT N. STILES
Department of Physiology and Biophysics, University
Health Sciences, Memphis, Tennessee 38163
SUMMARY AND CONCLUSIONS
1. Acceleration tremors of the extended
hand and surface EMGs of two extensor
muscles were detected simultaneously for
each of nine normal subjects. The raw, sur-
face EMGs were full-wave rectified and
smoothed, providing demodulated EMG
records. Auto- and cross-spectral analysis
of 16-s records of demodulated EMG and
hand acceleration was performed, giving
values of root mean square (rms) EMG at the
major tremor frequency and EMG-tremor
coherence. These two quantities were deter-
mined for steady-state displacement ampli-
tudes (rms values) of the tremor that ranged
between about 30 and 30,000 pm. Hand
tremors having this range of displacement
amplitudes were obtained during continuous
extension of the hand for lo-60 min, as
described previously (24).
2. The rms demodulated EMG amplitude
at the major tremor frequency and the EMG-
tremor coherence values were calculated
with the assumption that the grouped electri-
cal potentials at the tremor frequency re-
sulted from neural feedback within the
neuromuscular system controlling the hand.
This assumption is supported by previous
findings (24) that the frequencies of both the
tremor and these grouped potentials de-
creased together with mass loading of the
hand.
3. EMG-tremor coherence was generally
high for all but the smallest amplitude (less
than 100 pm) tremors. For these small-dis-
placement tremors, the EMG-tremor co-
herence decreased as both frequency and
displacement amplitude of the tremors de-
creased, suggesting a rate-sensitive mech-
anism in the neural feedback pathway.
40www.physiology.org/journal/jn
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of Tennessee Center for the
4. For each subject, the relation between
log(EMG) and log(displacement) was ap-
proximately linear, indicating that the de-
modulated EMG and the rms displacement
of the tremor are related by a power func-
tion. For the nine subjects, the tremor dis-
placement increased approximately as the
EMG squared. This relation is similar to that
found previously (28) for the soleus EMG
and ankle tremor for a similar three-decade
range of displacement amplitudes.
5. The power relation between EMG and
tremor displacement indicates that the ratio
of demodulated EMG to tremor displace-
ment generally decreased with increasing
displacement amplitude. A similarity is
suggested between the behavior of this ratio
and the sensitivity of primary endings of
muscle spindles of the decerebrate cat
(13, 18).
6. The effect of mass loading of the hand
on the tremor period appreared to depend on
the displacement amplitude of the tremor.
For small-displacement oscillations, mass
loading increased the tremor period, as ex-
pected for a constant-spring, variable-mass
system. For certain large-displacement
tremors (and certain subjects), the tremor
period was relatively independent of mass
loading. However, the period of large-dis-
placement tremors was found to depend on
hand position. Therefore, the period of these
oscillations may have varied with the
average muscle tension as well as with
neural feedback factors. These findings have
implications relative to those that suggest
an independence of either Parkinsonian
tremor of mass loading (25, 3 1) or ankle
clonus and Parkinsonian tremor of rhythmi-
cal forcing (30, 31).
Copyright
0 1980 The American(192.236.036.029)
${individualUser.surname} Physiological Society
on December 27, 201
 MECHANICAL-REFLEX
7. As reported previously (6), minor
bands between 9 and 15 Hz were sometimes
present in the acceleration spectra. These
bands occurred in addition to the major
band, which varied in frequency with either
mass loading or increased displacement
amplitude. These minor spectral bands were
found only occasionally for certain subjects,
and seemed to be absent when the major
tremor oscillation occurred at displacement
amplitudes above about 100 ,um.
8. The results of this study suggest that
internal perturbations, muscle-load me-
chanics, and neural feedback factors act
together to determine the many different
steady-state values of displacement ampli-
tude and frequency of hand tremor.
INTRODUCTION
Postural hand tremor in normal humans
is generally a rotational, involuntary oscil-
lation that occurs at a frequency between
about 8-10 Hz and at a very small dis-
placement amplitude (an rms angle of about
1’). While the mechanism of hand tremor is
not completely understood, the frequency
and amplitude of these oscillations appear to
be determined by a combination of neural
and mechanical factors of the particular
neuromuscular-load system. The finding
(24) that, under certain conditions, postural
hand tremor of normal man decreases in
frequency from about 9 to about 4 Hz as the
rms displacement amplitude increases from
about 30 to about 30,000 ,um is of particular
interest relative to the mechanism of these
oscillations. The similar quantitative rela-
tion (a negative-power rule) found between
the frequency and displacement amplitude
of hand and ankle tremor detected from
normal subjects (24, 28) and of hand tremor
from Parkinsonian subjects (25) provides
support for Eshner’s (7) suggestion in 1897
that all tremors of man may have a common
mechanism. Furthermore, the finding (25)
that both the frequency and rms displace-
ment amplitude of postural hand tremor
of both normal and abnormal (Parkinsonian)
man may depend on combined mechanical
and neural feedback factors has led to the
proposal of a mechanical-reflex oscillator
hypothesis (25) for the mechanism of these
oscillations. However, the mechanical-
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OSCILLATOR HYPOTHESIS 41
reflex oscillator hypothesis has been pro-
posed primarily as a modification of the
mechanical oscillator hypothesis, and has
yet to be considered in any detail.
The mechanical oscillator (mechanical
resonance) hypothesis for postural tremors
was described in some detail in 1967 (27)
for the human hand and finger, and in 1974
(21) for a muscle-load system of the rat. For
this hypothesis, a lightly damped visco-
elastic-mass system (muscle elasticity and
load inertia) that is continuously forced by
irregular disturbances (asynchronous con-
tractions of motor units) is considered
largely to account for both the frequency
and amplitude of the normally occurring,
small-displacement (less than 100 pm) hand
tremors. Under conditions of rest between
recordings, both the frequency and displace-
ment amplitude of postural hand tremor
have been found (24) to vary quantitatively
as predicted by such a lumped-parameter,
viscoelastic-mass system. However, the
frequency-displacement amplitude results
showed considerable variability between
subjects, suggesting that factors other than
load inertia were varying during these
experiments.
For the mechanical oscillator hypothesis,
the nervous system is thought to make two
major contributions to the tremor. First,
neural control of the mean position of the
limb may set the level of stiffness of an
equivalent elastic element of muscle acting
in series with the inertial load (27). In this
manner, the nervous system indirectly (and
partially) determines the resonant fre-
quency (the tremor frequency) of the muscle-
limb system. The second contribution is to
provide the continuous, but largely asyn-
chronous, contractions of the active motor
units. These asynchronous contractions in
turn act as small internal disturbances to
the lightly damped mechanical system and,
therefore, act (with other disturbances) to
maintain the small-amplitude tremors. For
this model, neural feedback (reflex) effects
are considered to play almost no role in
determining the frequency and only a minor
role (if any) in determining the amplitude
of the tremor (27).
While it appears that neural feedback does
little to determine either the frequency or
displacement amplitude of small-amplitude
 42
hand tremors, there is evidence for neural
feedback signals generated by these limb
oscillations (see Fig. 1; also see Fig. 7 of
Ref. 24). The presence of a band in the de-
modulated extensor EMG spectrum at the
tremor frequency, and the reduction of the
frequency of this band with mass added to
the hand (see Fig. 7 of Ref. 24), together
indicate that this electrical activity results
from neural feedback.
In attempting to determine the role of
neural feedback in the mechanism of hand
tremor, certain quantitative relations be-
tween the demodulated extensor EMG and
the tremor may be of interest. Estimates of
the phase between, and the relative ampli-
tudes of, the EMG and tremor for different
amplitudes and frequencies of the tremor
might suggest whether the neural feedback
is positive or negative (18, 19, 23). How-
ever, the importance of these estimates
would depend in part on the EMG-tremor
coherence (2). Recent evidence for postural
ankle tremor from normal subjects (21) in-
dicated that both the soleus EMG-tremor
coherence and the amplitude of the EMG
modulation at the tremor frequency were
very small for the smallest amplitude (below
about 20 pm) tremors. However, both
quantities increased with increased rms
displacement amplitude of the tremor above
this control level (see Fig. 5 of Ref. 2 l), sug-
gesting underlying motor-unit recruitment
and/or synchronization (5, 15- 17).
In general, little attempt has been made to
determine if any quantitative relationship
exists between the electrical activity of
skeletal muscle resulting from neural feed-
back and the postural hand tremor detected
from either normal or abnormal subjects.
The major purpose of this work is to report
the amplitude of the demodulated extensor
EMG activity at the hand tremor frequency
and the correlation (coherence) between the
EMG and tremor oscillations as the dis-
placement amplitude of the tremor increases
from control values of about 30 pm to values
of lOO- 1,000 times control. These values
are also considered for different amounts of
mass added to the subject’s hand. Finally,
the relation between the tremor period and
the logarithm (base 10) of the rms displace-
ment amplitude of these oscillations
reported.
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R. N. STILES
It is important to consider that a 9- to 12-
Hz oscillation of the hand may occur in ad-
dition to the hand tremor that depends on
mechanical factors (6). When present, this
9- to 12-Hz oscillation can be separated in
frequency from the mechanical tremor by
mass loading of the hand. An oscillation of
the forearm with similar frequency (9-12
Hz) has been found when that limb is either
mass (8) or spring (11) loaded. The appear-
ance of correlated electrical activity of the
muscles controlling these limbs suggests
either a reflex or central oscillator mech-
anism for these oscillations. While not
systematically studied, these 9- to 12-Hz
.oscillations of the hand seem to occur at
rms displacement amplitudes that are small
relative to the amplitude of the mechanical
tremor. In general, these oscillations (when
present) appear to be a minor component
of hand tremor.
METHODS
Hand tremor was detected from nine normal
human subjects with a small, sensitive ac-
celerometer while each subject held an unsup-
ported hand in an approximately horizontal
mean position. Each subject was seated with
the forearm pronated and supported to the wrist
in a horizontal position by a solid cradle mounted
on a short, heavy table. Two strips of 2-inch-
wide duct tape held the forearm tightly against
the cradle. The extended fingers and adducted
thumb were held tightly between two sheets of
plastic material (forming a sandwich, with added
mass of 100 g). Each subject was asked to keep
the mean position of the hand (determined
visually) approximately horizontal. Tremor in
the vertical plane was detected using an AVR-
250 accelerometer (2.5 g) taped to the upper
plastic sheet near the lateral-medial midline at
16 cm from the subject’s wrist. (The response
characteristics and method of calibration of this
accelerometer and associated electronics have
been described previously (26).)
In addition to hand acceleration, bipolar sur-
face EMG records from the extensor digitorum
communis (EDC) and extensor carpi radialis
longus (ECR) muscles were obtained simul-
taneously using Beckman bipotential electrodes.
Two electrodes were placed parallel with the
longitudinal direction of each muscle about 2 cm
apart, center to center. The center points of the
two pairs of electrodes were generally about 5 cm
is apart, and the two indifferent electrodes were
located on the lateral surface of the upper arm.
 MECHANICAL-REFLEX
Cross-correlation analysis of the two simulta-
neously obtained EMGs indicated that the sur-
face potentials detected by one set of electrodes
were not detected by the second set. Amplifica-
tion was obtained using Tektronix (model 122)
preamplifiers (bandwidth of 8- 1,000 Hz) and
locally constructed DC amplifiers. Any motion
artifact in the EMG signals was eliminated by
the combined use of the ~-HZ high-pass filter
setting on the preamplifiers and an additional
high-pass digital filter. The power spectral
density of the noise of the amplifiers did not
exceed 1 pV*/bandwidth for any part of the
spectrum.
Hand tremor and extensor EMG records (16-s
records) were obtained for analysis under five
different conditions.
I) Rest between records. Different 16-s ac-
celeration records were obtained with approxi-
mately 90-s rest between each pair of runs.
During the rest period, the subject relaxed the
muscles controlling the hand, allowing the hand
to rest in a nearly vertical position. From 5 to
20 records with rest before each 16-s run were
obtained for analysis from each subject studied.
2) Different amounts of added mass and rest
between records. Four sets of 30 tremor records
each were obtained from three subjects under
conditions of different amounts of mass (lead
weights of 0, 100, 200, 300,400, and 500 g) taped
tightly to the ventral part of the sandwich at
approximately 14 cm from the wrist. The order of
addition of these weights was determined from
a random-number table. An approximately 90-s
rest period was taken before each recording
period.
3) No rest between records. For these ex-
periments, each of the nine subjects maintained
the hand (the right hand) in a mean horizontal
position without rest for a total period of between
10 and 60 min. (This length of time of con-
tinuous maintenance of a mean horizontal posi-
tion of the hand led to little or no discomfort
for any of the subjects.) This procedure generally
resulted in an increase in the tremor displace-
ment amplitude over time (see Fig. 3).
During most of each lo- to 60-min period of
continuous extension of the hand, 16-s digitized
records were obtained every l-5 min. However,
no attempt was made to obtain a fixed num-
ber of recordings at each session, nor to
obtain records with a fixed time period be-
tween them. In general, an attempt was made to
obtain 16-s records somewhat uniformly through-
out the different steady-state amplitude levels as
the tremor increased from very small levels
(about 30 pm) to a level between about 0.5 and
3 cm. It is important to note that the rms tremor
amplitude (acceleration amplitude) generally in-
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OSCILLATOR HYPOTHESIS 43
creased very little (if any) with time through-
out any given 16-s digitization period.
This sequence of recordings without rest was
obtained from three of these subjects with dif-
ferent amounts of mass added to the hand. In
addition, this sequence was obtained twice for
subject I for each of 100 and 500 g, and 3 times for
subject 5 for each of 100 and 300 g.
For one subject whose tremor displacement
amplitude increased very little over a 60-min
period, a second sequence (session 2) of record-
ings was made under the same conditions (no rest
and 100 g added mass) approximately 1 mo after
the first.
4) Different mean positions of the hand. For
one subject, postural hand tremor was detected
for three different mean positions of the hand.
For these three sets of runs, the subject main-
tained the mean position of the hand at approxi-
mately 30” below the horizontal for the first set,
horizontal for the second, and 30” above the
horizontal for the third. For each set, postural
tremor was detected with and without rest of the
hand between runs.
Each 16-s record of the voltage analog of hand
tremor acceleration was digitized by a PDP-12
computer at a rate of 64/s (a rate sufficient to
ensure no aliased peaks in the power spectra).
The mean was computed and subtracted from
these 1,024 digital values before they were stored
on magnetic tape for subsequent auto- and cross-
spectral analysis. Sixteen-second records of the
EMGs were digitized at 1,024/s, estimates of the
means were computed and subtracted from the
data, and then these deviations were full-wave
rectified and smoothed (16 nonoverlapping
values were averaged). This process results in
amplitude demodulation of the EMG signals
and gives an effective sampling rate of 64/s.
For the analysis of these tremor and EMG
data, auto- and cross-covariance functions were
computed with 64 and 128 (+64) lags, respec-
tively. Auto- and cross-spectral functions were
then computed from these covariance functions.
Each spectrum included 65 frequency values
from 0 to 32 Hz at intervals of 0.5 Hz (a band-
width of 0.5 Hz). Each EMG spectrum was cor-
rected for the attentuation of the amplitude as a
function of frequency caused by the digital filter.
In general, each tremor spectrum (power, or
variance, spectrum) obtained in this study con-
tained primarily one major spectral band, a band
with a peak frequency, f, that was determined
to the nearest 0.25 Hz (see Ref. 24 for a more
detailed description of this analysis).
Finally, coherence values, rs,‘(f), were com-
puted from the combined auto- and cross-
spectral values. As computed, ysJ,*(f) is an esti-
mate of what percentage of the variation of&) is
 44 R. N. STILES

correlated with variation of y(t) at the frequency

average rms displacement was 25 pm for
f Hz. The coherence will be zero if x(t) and y(t)the first set (Fig. IA) and 66 pm (Fig. 1B)
are statistically independent, or unity if x(t) for the second. In comparing the two
and y(t) are completely correlated. Coherence averaged EMG spectra of Fig. 1, note that
values between zero and unity indicate either that
the total variance of the band at the tremor
other factors (perhaps noise) are determining one
of the variables but not the other, or that the frequency is slightly higher in B than in A.
system relating x(t) and y(t) is not linear (2).Also note that there was a larger average
coherence between the EDC EMG and
For the number of digital values (samples) ob-
tained for each 16-s run (1,024 samples) and thetremor associated with the larger displace-
number of quadratically smoothed spectral esti- ment amplitude tremor.
mates (64 spectral values), each smoothed From Fig. 1, it is of interest to note the
spectral value had 32 deg of freedom. For this presence in A of the prominent band with
number of degrees of freedom, the minimum peak frequency at approximately 19 Hz, and
EMG-tremor coherence value that can be taken its absence in B. First, this band did not
to be different from zero at the 95% level of con-
result from harmonic distortion of the raw
fidence is about 0.20. However, all the coherence
values calculated for each subject were plotted EMG signal due to saturation of the ampli-
in this report, even though some values were notfiers or the A/D converter of the computer.
While not studied in detail, it appeared from
statistically significant. For the acceleration and
visual inspection of the raw EMG records
EMG spectra, a peak-to-valley ratio of 2 to 1 was
necessary for a spectral value to be significant that this band at 19 Hz was due to the de-
at the 95% level of confidence. tection by the surface electrodes of regular,
large-amplitude potentials, perhaps due to
RESULTS the activity of a single motor unit. It may
be that the absence of this band in Fig. 1B
Presence of a band in EMG spectrum at resulted from a slightly different placement
major frequency of hand tremor of the electrodes on the days that these runs
Associated with most of the postural were taken. However, bands between 12
tremor records that were obtained with rest and 20 Hz were present in the EMG spectra
between runs and 100 g added mass was a for others of the nine subjects, and generally
modulation of the amplitude of the EDC and disappeared as the displacement amplitude
ECR EMG records at approximately 8 Hz. of the subject’s tremor increased.
The rms amplitude of this EMG modulation
(when present) was very small (generally Presence of bands in EMG spectrum
between l- 10 pV). While a few of the correlated with minor bands in
EMG spectra associated with 16-s records acceleration spectrum
of small displacement tremors did not show For each of the nine subjects, bands with
a statistically significant band at the tremor peak frequencies between 9 and 20 Hz ap-
frequency, averaging a number of these peared in certain individual EDC and ECR
spectra generally revealed a highly signifi- spectra. However, for seven of the nine
cant band at this frequency. Figure IA andB subjects, correlated bands appeared in a few
illustrates two such average EMG spectra, of the associated acceleration spectra at
each of which contains a band at the frequencies between about 9 and 15 Hz, as
average tremor frequency (about 8 Hz). shown in Fig. 2 for three subjects. For two
These two plots illustrate an average of of these seven subjects (subjects I and 2),
20 (A) and 10 (B) EMG spectra for EDC a band with peak frequency of about 10 Hz
EMG records obtained from subject 2. appeared in the acceleration spectra when
These two sets were obtained on two differ- the hand was mass loaded (200400 g) (see
ent days (about 1 yr apart). Each set of runs Fig. 2A). For one subject (subject 3), a band
was obtained with 100 g (the two plastic at 1 l- 12 Hz appeared in a few acceleration
sheets) added to the hand, and with rest be- spectra, but only when the subject’s hand
tween runs of the muscles controlling the was not mass loaded (0 g added mass). (This
hand. While the average tremor frequency band at 11-12 Hz appeared in addition to
was nearly the same for both sets of data, the the major band at approximately 9 Hz.) With

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 MECHANICAL-REFLEX OSCILLATOR HYPOTHESIS

A Subj. I N= 20 B Subj. I N = IO
olsp = 25 pm Dlsp = 66 pm
7 = 8.3 Hz i = 8.2 Hz
-2 = 0.55 ;2 =
Y 0.70

FREQUENCY (Hz)
FIG. 1. Each graph (A and B) shows averaged EMG spectra of demodulated EDC records obtained from
subject I. These records (20 and 10 for spectra A and B, respectively) were obtained with 100 g added mass
and rest of the hand between recordings. Average tremor displacement, frequency, and EMG-tremor
coherence values are shown for each set of records. Each arrow denotes the average tremor frequency.

100 g added to this subject’s hand, the ll- subject maintained the hand at a mean
to 12-Hz band was absent, and only a single horizontal position for a period of time
band with peak frequency between 7.5 and without rest.
8.25 Hz appeared in the acceleration For a few subjects, it is possible that the
spectra. Therefore, for two subjects, a major and minor bands may have merged for
minor band with peak frequency at about certain runs. However, there is no evidence
10 Hz was present in the tremor (and EMG) to indicate that this occurred. The minor
spectra only with the addition of a certain band, if absent, was never uncovered in the
mass load, while a similar band disappeared spectrum when the frequency of the major
with mass loading for a third subject. In band decreased with increased displace-
contrast, the acceleration spectra of a fourth ment amplitude. Also, for those sessions
subject (subject 5) whose hand was mass with at least 100 g added to the subject’s
loaded (added masses of 0, 100, and 300 g) hand, the frequency of the mechanical-
showed no significant minor bands between reflex tremor was generally below the 9-
9 and 15 Hz. to 15-Hz range of frequencies within which
In general, the rms displacement ampli- the minor band appeared.
tude of the 9- to 15Hz hand oscillations In Fig. 2, the coherence values between
was between 10 and 20 pm (see Fig. 2). For the EMG and acceleration bands are shown
those experiments in which the mass of the next to the arrows in the EDC and ECR
subject’s hand was varied, only the promi- EMG spectra. For certain of the minor
nent band changed systematically in fre- bands (between 9 and 15 Hz), a highly co-
quency with the added mass. In addition, herent band appeared in both the EDC and
this same major band was the only one that ECR EMG spectra (note the 10.25-Hz band
increased in displacement amplitude as the in Fig. 2C). In contrast, a correlated EMG

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 46 R. N. STILES

A Subj. I B Subj. 8 C Subj. 4

+500 gm +I00 gm + 100 gm

TREMOR

61 pm 93 pm 32 Pm
\r ‘; (8.5 Hz) ’ - (8.25 Hz)
I .0
‘i (6*5 Hz) I’ ?I
!I I’ I I 17 pm

)- 0.4
I-
z 0.2
z 8
w 0 1
n 0’ B

-J
a EDC EMG
a / 0.08 ,0.50
I-
O
w
L
w

a
w
9
0
e

W
N 0’
Ts ECR EMG
a ,0.45 ,0.28 ;0.54
E
a
0
I.0
ll
rt l
‘\
‘\
00
* 0.8 ’ \ 1 \
Ie
I I

006 0.23 ;
t L I

Oh
0 5 IO I5 20 0 5 IO 15 20

FREQUENCY (Hz)
FIG. 2. A-C: plots of single, normalized spectra of hand acceleration, EDC EMG, and ECR EMG
records for each of three subjects (subjects I, 7, 4). Numbers near each arrow in the tremor
spectra denote the peak frequency and rms displacement for each band. Each number in the EMG spectra
denotes the EMG-tremor coherence at the peak frequency of the corresponding acceleration band.

band would at times appear in only one of 2B). Since the main interest in this study was
these two EMG spectra (see the 11.5Hz the band in the acceleration spectrum that
band in the acceleration spectrum of Fig. decreased in frequency with added mass,

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 MECHANICAL-REFLEX
these additional oscillations of the hand (at
higher frequencies and smaller displacement
amplitudes) were not studied further.
EMG amplitude, EMG-tremor coherence,
tremor period, and tremor displacement
amplitude for different sequential
digitization periods (runs)
For most subjects, the tremor displace-
ment amplitude increased with time as the
subject extended the hand horizontally
without rest against the force of gravity.
Other quantities that generally increased
with tremor displacement amplitude were
the rms extensor EMG amplitude (in pV)
determined at the major tremor frequency,
the EMG-tremor coherence, and the tremor
period.
The variations of these four quantities
during a sequence of digitization runs are
shown in Fig. 3A-C for three different
sequences (two subjects). From bottom to
top, the logarithm (base 10) of the rms
tremor displacement (in pm), the coherence
between the EDC EMG and tremor, the
tremor period (in ms), and the logarithm
(base 10) of the rms EMG amplitude (in pV)
at the tremor frequency are plotted against
run number. In general, these values were
obtained for an initial sequence of 10 runs
that occurred with rest of the hand between
runs, followed by a sequence with no rest,
and ended with a second sequence of runs
with rest.
Figure 3A and B shows these values
plotted against run number for subject 1 with
100 and 500 g added to the hand, respec-
tively. Note that the levels of the log(EMG)
are approximately the same in A and B for
similar log(displacement) levels. Also note
that the tremor period appears to be gener-
ally higher in B than in A. Also, it is evident
from Fig. 3B that as the log(displacement)
begins to increase above control values
(values obtained with rest), the tremor
period decreased initially before increasing
with the tremor displacement amplitude.
Note that the EMG-tremor coherence asso-
ciated with the small-displacement tremors
is relatively small (approximately 0.2) for
the runs obtained with the 500-g mass added
to this subject’s hand. However, this co-
herence increased to nearly unity under
both conditions of added mass (100 and
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OSCILLATOR HYPOTHESIS 47
500 g) for tremors having displacement
levels above about 1,000 ,um.
Figure 3C shows these four quantities
plotted against run number for subject 2,
with 100 g added mass. However, in con-
trst to the protocol for subject I, this sub-
ject rested the hand for 2 min after run 22,
and then began another period of continuous
extension of the hand. This second no-rest
period continued through run 33, after
which the subject again rested the hand be-
fore each subsequent run. Note that, associ-
ated with the decreased tremor displace-
ment that occurred after the 2-min period of
rest between runs 22 and 23, the other three
quantities all decreased approximately to
control levels.
For about half of the recording sessions
in this study, there was a relatively gradual
increase in tremor displacement amplitude
with time during continuous extension of the
hand (see Fig. 3A and B). However, for
certain subjects (on certain days), a gradual
increase in amplitude occurred initially, fol-
lowed by a rapid increase to a much larger
level (see Fig. 3C). Also, for certain sub-
jects, the displacement amplitude increased
to relatively large levels (1 ,OOO- 10,000 pm)
and then decreased spontaneously (abruptly,
during continuous extension of the hand) to
near control levels (100-500 pm), followed
again by an increase to large displacement
levels. For certain sessions (and subjects),
maximum displacement amplitude (greater
than 10,000 pm) was reached after about
10 min of continuous extension of the hand.
In contrast, the displacement amplitude of
the tremor of subject 6 failed to reach 1,000
,um after 60 min without rest on two separate
occasions.
Tremor period and rms
displacement amplitude
The plots of Fig. 3A and C indicate that
the tremor period increased to slightly
greater than double the control level as the
tremor displacement amplitude increased
nearly l,OOO-fold. Also, the period of the
small-displacement tremors (less than 100
,um) increased by about 50 ms with 500 g
of mass added to the hand (compare the
triangles of Fig. 3A and B). A more quanti-
tative comparison between tremor period
and log(displacement) values for five of the
nine subjects (and for different added
 48 R. N. STILES

A Subject I B Subject I C Subject 2

+ 100 gm + 500 gm + 100 gm

1,
I
I
A# f
27
Al+
I
250 AA’
I ’
AI
225 A I
I
200 % I
’ I

0.8

0.6

0.4
i=
0.2 E
E
0 2 Ii II
2 ;I Il
ov> 11 Il
Q II ll

REST RE%T No REST

,- --REST---
xl l ’ 1 I I
40 0 IO 20 30 40 0 IO 20 30 40
-NO--
REST
t
2 MIN REST
RUN NUMBER
FIG. 3. A-C: each of the four sets of points shows (top to bottom) log(rms EMG), tremor period,
coherence, and log(rms displacement) values plotted against run number for: A, subject I with 100 g;
B, subject I with 500 g; and C, subject 2 with 100 g added mass. These values were calculated for both
rest and no rest of the hand between runs.

masses) is shown in Fig. 4A -D. For all nine range of log(displacement) values (see Fig.
subjects with 100 g added mass, the relation 4A and B) or to be similar to that shown in
between these two variables appeared either Fig. 4C and D for subjects 3 and 4. For
to be approximately linear over most of the subjects 3 and 4, note that the tremor

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 MECHANICAL-REFLEX OSCILLATOR HYPOTHESIS 49

A Subj. I C Subj. 3
0 100 gm 0 Ogm
0 500 gm l IOOgm
L
t

B Subj. 5 0 l Subj. 2 (100 gm)

A Ogm 0 Subj. 4 (100 gm)
w I 0 100 gm

E t
0 300 gm
I
225

125

I I I
75’
I 2 3 4

LOG (RMS DISPLACEMENT)

FIG. 4. A-D: tremor period plotted against log(rms displacement) for subjects Z-5 for different masses
added to the hand. A, for subject I with 100 and 500 g; 23, for subject 5 with 100 and 300 g; C, for subject 3 with
0 and 100 g; and D, for subjects 2 and 4 with 100 g. Each line was drawn to illustrate the apparent relation
between tremor period and log(displacement) values.

period changed very little or decreased interest that each set of points shown in Fig.
slightly with increasing displacement ampli- 4A for subject 2 represents values obtained
tude for values below about 1,000 ,ccm. (The on two separate occasions. Since the plots
lines in Fig. 4 are not regression lines, but were very similar for each occasion with the
rather are drawn to emphasize the apparent same added mass, no distinction was made
relation of the data.) on this basis in plotting either set of values.
The effect of mass on the relation between The data for subject 5 in Fig. 4B are of
tremor period and log(displacement) ap- interest because they indicate that in-
peared to depend on both the mass and the creasing the mass of this subject’s hand by
subject. For subjects 2 and 3 (Fig. 4C and 300 g increased the period for only the small-
D), the tremor period-log(displacement) and large-displacement amplitude tremors.
plot was shifted systematically toward This behavior appeared to be characteristic
greater values of period by additional mass for subject 5, as determined from plots (not
added to the hand. While not illustrated, an illustrated) of paired values obtained on two
additional set of points obtained for subject additional occasions for each of 100 and
1 with 300 g added mass fell largely between 300 g added mass. With 300 g, the tremor
the two lines drawn in Fig. 4A. Also, it is of period-log(displacement) relation appeared

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 50 R. N. STILES

to be U-shaped (similar to the plot in Fig. that the magnitude of the EMG-tremor
4B) for each additional set of data. For coherence may be reduced in some manner
100 g, the additional two plots were similar by mass added to the hand.
to the one shown in Fig. 40 (open circles), Plots of the coherence between the
with the exception that seven additional demodulated EDC EMG and hand tremor
paired values were obtained at approx- for different log(displacement) levels of the
imately 10,000 pm and 210 ms. For either tremor are shown in Fig. 5 for five of the
100 or 300 g added mass and tremor nine subjects. The plots for three of the
displacement amplitudes between about remaining subjects with 100 g added mass
10,000 and 30,000 pm, the tremor period were similar to that shown in Fig. 5C for
ranged between 150 and 235 ms for this subject3 with 100 g added mass. In general,
subject. the coherence-log(displacement) plots were
similar for both the EDC and ECR EMGs,
EMG-tremor coherence and tremor for the same subject and amount of added
displacement amplitude mass. The points plotted in Fig. 5A (sub-
The plots of Fig. 3 indicated that the EMG- ject I) are of interest in that each set repre-
tremor coherence was statistically significant sents values obtained on two different
for most of these tremor recordings, and occasions with the same amount of added
increased either slightly (Fig. 3A) or consid- mass (either 100 or 500 g). However, be-
erably (Fig. 3B) with the log(displacement) cause of the similarity of the plots obtained
amplitude of the tremor. A comparison of the for the same added mass, no distinction
coherence values of Fig. 3A and B suggests was made in Fig. 5A between these sets.

A Subj. I 0 100 gm C Subj 3 oogm

0 500 gm 0 100 gm
--------------- ------e--m----

1.0

0.8

0.6

0.4

Nk 0.2

0
w -
Y B Subj. 5 0 100 gm D l Subj. 2 (IO0 gm)
----------------- 0 300 gm 0 Subj. 4 (IO0 gm)
w-B------------

LOG (RMS DISPLACEMENT)

FIG. 5. A-D: EMG-tremor coherence (EDC EMG) at the major frequency of hand tremor plotted
against log(rms displacement) for subjects I-5 for different added masses. A, for subject I with 100 and
500 g; B, for subject 5 with 100 and 300 g; C, for subject 3 with 0 and 100 g; and D, for subjects 2 and 4 with
100 g. Each line was drawn to illustrate the apparent relation between coherence and log(displacement).

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 MECHANICAL-REFLEX
Also, an additional set of points was ob-
tained for subject 5, with 0 g added mass.
However, the plot of this set of values was
very similar to the two sets shown in Fig.
5B for 100 and 300 g, and is therefore not
included in this figure. (The plots obtained
for 100 and 300 g on two additional oc-
casions for subject 5 were also very similar
to plots shown in Fig. 93.)
For eight of the nine subjects (all except
subject 6) with 100 g added mass, the EMG-
tremor coherence increased from values of
about 0.2 to values between 0.6 and 0.8 as
the rms displacement of the tremor in-
creased from about 30 ,um to a level of about
100 ,um. Between 100 ,um and the maximum.
value (lO,OOO-30,000 pm), the coherence
generally increased gradually from 0.6 to 0.8
to a maximum of very near unity. The
tremor of the remaining subject’s hand (sub-
ject 6) reached a maximum of only 500 ,um,
even though the experiment was continued
for 60 min on each of two separate sessions.
For a total of 66 runs for this subject, the
EMG-tremor coherence values were all
relatively high (between 0.54 and 1.1) as
the tremor displacement amplitude varied
between 50 and 500 ,ccm.
The effect of different added masses on
the coherence-log(displacement) relation
appeared to differ somewhat depending on
the subject and the magnitude of the mass,
as shown in Fig. 5A-D. However, when the
coherence values obtained for a single sub-
ject with different added masses were
plotted (not illustrated) against the logarithm
of the rms acceleration of the tremor, there
appeared to be little difference between
sets of data. For subjects I and 3 (Fig. 5A
and C), the open circles were shifted to the
right and approximately overlay the filled
circles. For subject 5, there was a slight
separation of the two sets of points shown
in Fig. 5B for coherence values below about
0.30. In general, the variability between
plots for different subjects with 100 g
appeared to be reduced for coherence-
log(acceleration) as compared with co-
herence-log(displacement) values.
EMG-tremor coherence and
tr’emor frequency
The data of Fig. 5A and C indicate that,
for certain subjects, increasing the mass of
the hand may lead to a reduced EMG-tremor
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OSCILLATOR HYPOTHESIS 51
+lOO-600 gm, WITH REST
+I00 gm, WITHOUT REST
FREQUENCY (Hz)
FIG. 6. Mean EMG-tremor coherence (EDC EMG)
values plotted for different tremor frequencies
between 4.25 and 9.25 Hz; for rest and different
added masses (filled circles), and for no rest and 100 g
added mass (open circles). Each line was drawn to
illustrate the apparent behavior of one set of points.
coherence for small-displacement amplitude
tremors. While an increased mass of the
hand may i.ndirectly lead to a small increase
in the displ acement amp1itude of th .e tremor
detected with rest between runs (perhaps
because of a decreased equivalent damping
ratio of the system), the added ma .ss con-
sistently led to a reduced tremor fre quency
under these conditions. Therefore, the re-
duced coherence with increased mass seen
in Fig. 5A and C may have resulted
from a decrease in the frequency of these
oscillations. In Fig. 6, the EMG-tremor
coherence (EDC EMG) has been plotted
against tremor frequency for two different
groups of experiments. For one group (filled
circles), each subject’s hand (three subjects
with a total of four sets of data) was rested
between run s, and EMG-tremor coherence
values were obtained for different amounts
of added mass (100-600 g). For the second
group (open circles), each subject’s hand
trem or (nine s bjects) was detec ted without
rest between runs and with a constant
amount of added mass (100 g). In this figure,
each circle represents the mean coherence
value calculated at a particular frequency.
(Each point represents a mean of about
seven values. )
For runs obtained with rest of the mu .scles
controlling the hand, an in creased mass
generally res ulted in an increased mean
displacem .ent amp1 tude, increasing from a
minimum of about 30 pm to a maximum
 52 R. N. STILES

of about 80 pm. Even though the tremor
displacement amplitude generally increased
with mass under these conditions, the
average EMG-tremor coherence decreased
from about 0.6 to values below the level
of significance (below 0.2). A plot of the
mean EMG-tremor coherence against
tremor frequency shows a positive, ap-
proximately linear, relation (filled circles
of Fig. 6) between these two quantities.
In contrast to the filled circles, each
open circle of Fig. 6 is an average coherence
value calculated from records obtained
without rest and with constant added mass
(100 g). These open circles of Fig. 6 il-
lustrate that, under these conditions, the
mean EMG-tremor coherence generally in-
creased as the tremor frequency decreased
over the same range found for the filled
circles. However, for these experiments,
the displacement amplitude increased from
very small values (tremors not visible to
the eye) to rms levels of about 3 cm (for
eight of the nine subjects).
Extensor EMG modulation at tremor
frequency relative to displacement
amplitude of tremor
From Fig. 3, it appears that the EMG
modulation (in pV) at the tremor frequency
increases rather systematically with the dis-
placement amplitude of hand tremor. The
similarity of the variation of the logarithm of
these two quantities with run number can be
seen from the three sets of data plotted in
Fig. 3. In Fig. 7A and B, the log(rms EMG)
has been plotted against the log(rms dis-
placement) of the tremor for all nine sub-
jects. Each set of points (for each subject)
is made up of paired values calculated from
EDC EMG and tremor records obtained
for runs with and without rest and with
100 g added mass. The lines in Fig. 7A were
drawn to illustrate the apparent behavior
of these points for subjects 1,5, and 7.
For each of six subjects (Fig. 7B), the
relation between log(EMG) and log(dis-
placement) was fairly linear, with a slope
similar to that of the other five sets. From

. B

18
.

G I.6

g I.0 l Subj. 2
A Subj. 3
-J 0.8 l Subj. 4
06
. 0 Sub] ‘.
6
o Subj. 8
04
. A Subj. 9

LOG (RMS D I SPLACEMENT)

FIG. 7. A and B: log(rms EMG) (EDC EMG) plotted against log(rms displacement) for nine subjects. A,
paired values for subjects I, 5, and 7, with lines drawn to illustrate the apparent behavior of the points; B,
paired values (six subjects) for which regression lines were calculated (see text).

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 MECHANICAL-REFLEX
calculation of the regression equations, the
slopes for these six sets of points were 0.35,
0.38, 0.46, 0.47, 0.50, and 0.51, while the
correlation coefficients ranged from 0.91 to
0.98. In contrast, the paired log values for
each of the remaining three subjects (Fig.
7A) appeared to follow two different linear
relations. Note that for each of these sets of
data, the slope of the relation appears to
change sharply at a displacement level of
about 100 ,um. Also note that two of these
sets are nearly identical in form and in mag-
nitude of the ordinate values. While the third
set (for subject 7) was similar in form to
these two, the EMG values are smaller at
similar displacement levels by a factor of
about one-fourth.
In addition to the plots of Fig. 7A and B,
which illustrate the relation between the
paired values of EDC EMG and tremor dis-
placement, plots were also made for each of
these nine subjects using the ECR EMG
values. For a given subject, the log(EMG)
and log(displacement) plots for these two
extensor muscles differed primarily in ab-
solute value of the ordinate (the log(EMG))
and very little in general form. For sub-
ject I, the rms EMG amplitude of the EDC
muscle was about double that of the ECR for
all tremor displacement amplitudes. How-
ever, for other subjects, the rms amplitude
of the ECR EMG either differed very little
from, or was slightly greater than, that of
the simultaneously obtained EDC EMG.
Ejjkct oj’mean position oj’hand on
tremor and correlated EMG activity
For subject 1, a sequence of hand tremor
and EMG records was obtained on the same
day for each of three different mean positions
of the hand. While not illustrated, the co-
herence, tremor period, and log(EMG)
varied with the log(displacement) in a man-
ner that was very similar for all three sets of
data. Also, these plots were very similar in
general form to those found for this subject
on other days (see the data shown in Figs.
4, 5, and 7 for this subject). In comparing
absolute magnitudes, the smallest rms dis-
placement and highest frequency values
(12 pm at 8.75 Hz), as well as the lowest
frequency (3.0 Hz with 3 1,OOO-pm displace-
ment),were all obtained with the hand held
30” below the horizontal. The smallest EMG
value (2 ,uV) was also found for this hand
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OSCILLATOR HYPOTHESIS 53
position, while the largest EMG value (280
,uV associated with an rms displacement of
35,000 pm) was obtained with the hand held
30” above the horizontal. These results
primarily indicate that large-displacement
tremors can be induced at different mean
positions of the hand. They also indicate
that the minimum and maximum values of
certain measures of the tremor and related
EMG are dependent on hand position.
For each of the three sets of experiments
described above, the mean position of the
hand was maintained approximately con-
stant throughout each sequence of runs.
However, it was found that, once large-
amplitude tremors were induced (after
continuous extension of the hand at one
mean position), they generally continued
as this subject moved the hand from one
mean position to the next. Figure 8 shows
hand tremor and demodulated extensor
EMG records obtained as the subject main-
tained the hand at three different mean
positions during a 16-s run. (These rec-
ords were obtained for illustration from
stored digital values.) As the mean position
of the hand changed from below to above
the horizontal, the frequency of this large-
amplitude tremor changed from a minimum
of 3.0 Hz (at 30’ below the horizontal), to
4.5 Hz (horizontal), and finally to 5.75 Hz
(at 30” above the horizontal). The approxi-
mate doubling of peak acceleration ampli-
tude associated with the increase in tremor
frequency from 3.0 to 5.75 Hz indicates
that the displacement amplitude decreased
with increased frequency of the tremor. It
should be noted that the EDC and ECR
EMGs are modulated at the tremor fre-
quency during each of these three positions.
While the rms amplitude of the modulation
is greater in the EDC than in the ECR EMG
(typical for this subject), this amplitude
modulation is visually more evident in the
ECR record.
DISCUSSION
The major purpose of this work was to
quantify both the amplitude modulation
of the surface EMG (EDC and ECR muscles)
at the major frequency of hand tremor and
the correlation (coherence) between this
EMG modulation and tremor in normal
subjects. These quantities were calculated

TREMOR
3.0 Hz
EDC EMG
ECR EMG
FIG. 8. Hand tremor (top), demodulated
records obtained simultaneously
amplitude tremor occurred at 3.0,4.5,
the horizontal, horizontal, and 30” above the horizontal,
for steady-state (rms) displacement ampli-
tudes of the tremor that ranged between
about 30 and 30,000 ,um. Auto- and cross-
spectral calculations were made with the
assumption that these grouped electrical
potentials at the major tremor frequency
resulted from neural feedback within the
neuromuscular system controlling the hand.
This assumption is strongly supported by
previous findings (24) that the frequencies
of both the tremor and these grouped
potentials decreased together with mass
loading of the hand.
While the grouping of muscle potentials
in the surface EMG at the major frequency
of hand tremor is of interest, Taylor (29)
has shown that such a grouping could occur
from chance synchronization of motor-unit
potentials. Statistically significant grouping
was found to be present (as indicated by
a significant band in the EMG spectrum;
see Fig. 1) in all but a few of the demodu-
lated EMG records obtained with 100 g
added to the subject’s hand. Also, with this
amount of mass loading, the EMG-tremor
coherence was generally high for all but
certain of the smallest displacement tremors
(see Fig. 5).
The significant grouping of potentials in
the EMG suggests some kind of underlying
motor-unit synchronization at the tremor
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R. N. STILES
4.5 Hz 5.75 Hz
360 /IV
180 pV
EDC EMG (middle), and demodulated ECR EMG (bottom)
for three different mean positions of the hand. Large-displacement
and 5.75 Hz as the mean position of the hand was maintained at 30” below
respectively.
frequency. As shown by Mori (16), motor
units having firing rates near (or at) the
tremor frequency might fire in synchrony
with the tremor. Also, motor units firing at
higher rates may have these ongoing rates
modulated at the tremor frequency in
synchrony with the slower tremor oscilla-
tion (5). Finally, motor units that normally
fire at ongoing rates higher than the tremor
frequency may be recruited to fire only once
each tremor cycle (some may fire twice,
producing doublets (4, 5)). This kind of syn-
chronization would be more likely to occur
during large-displacement tremors and also
be more likely to involve high-threshold
(large) motor units (4). Such units, when
recruited to fire only once per tremor cycle,
would appear to have an ongoing (tonic)
firing rate equal to the tremor frequency.
While a different neural mechanism could
account for each kind of synchronization,
all three kinds could occur at the receptor
level (perhaps muscle spindles) in the neural
feedback pathway, i.e., the tremor oscilla-
tion could be causing the synchronization,
rather than the converse. In a closed-loop
system with many feedback pathways (and
many inputs), it is difficult to make in-
ferences about cause and effect. Therefore,
a conclusion that “grouped discharges of
many motor units are the neuronal me-
 MECHANICAL-REFLEX
chanism of tremor” (4) should be considered
very carefully.
In this study, coherence values fell below
the level of statistical significance (below
0.20) only for certain hand tremor records
with either small displacements or low fre-
quencies (resulting from mass loading) com-
bined with small displacements. The de-
crease in EMG-tremor coherence with de-
creased frequency for small-displacement
tremors (the solid circles of Fig. 6) strongly
suggests that the feedback at the tremor
frequency may be proportional to the time
rate of change (velocity or acceleration)
of the oscillatory motion.
In considering the mechanical-reflex
oscillator hypothesis for hand tremor, it
appears that muscle-load mechanics largely
determine the frequency (24,27) and at least
partially determine the amplitude (24) of the
small-displacement (below 100 pm) oscilla-
tions. Even though the EMG-tremor co-
herence varied between very small values
(values below the level of significance) and
rather high values (0.6-0.8) for these small-
displacement hand tremors (for certain runs
and subjects), the tremor frequency changed
very little at these amplitude levels with
constant mass load. It appears that the
equivalent elasticity (muscle and reflex) of
the system remained relatively constant
for these tremors. The equivalent elasticity
also appeared to remain relatively constant
as the tremor frequency varied systemati-
cally with mass loading (27). Under these
conditions, the EMG-tremor coherence de-
creased with frequency from rather high
values to values below the level of sig-
nificance (Fig. 6). This large variation in
EMG-tremor coherence with apparent con-
stancy of the equivalent elasticity suggests
that a mechanical oscillator mechanism is
largely determining the frequency of these
small-displacement oscillations (27).
If mechanical factors (muscle elasticity
and load inertia) primarily determine the
frequency of small-displacement hand
tremors, then the neural feedback may
primarily affect the amplitude of these oscil-
lations. While it has been suggested (27)
that neural feedback may affect the tremor
amplitude by adding to the damping of the
system, this effect is apparently very small
for small-amplitude tremors. This is shown
by findings (20, 27) that the equivalent
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OSCILLATOR HYPOTHESIS 55
damping ratio for the neuromuscular-hand
system is normally between about 0.1 and
0.2. Certainly a major problem in deter-
mining the effect of neural feedback on
tremor amplitude is that the muscle-load
mechanics also contributes to the system’s
damping ratio (21). Another problem is that
the amplitude of the tremor will be af-
fected by the spectral characteristics of the
internal perturbations (asynchronous motor-
unit contractions, the pressure pulse).
Therefore, the displacement amplitude of
these small tremors might not vary con-
sistently with any particular measure of
neural feedback, e.g., EMG amplitude or
EMG-tremor coherence. Certainly the ordi-
nate values of Fig. 5 (EMG-tremor co-
herence) and Fig. 7A (log(EMG)) for small-
displacement tremors illustrate consider-
able variation in these quantities with
very little change in the rms displace-
ment amplitude. However, the data of Fig.
7B for six of the nine subjects show a rather
systematic (approximately linear) increase
in log(EMG) values with increasing log(dis-
placement) for these small-amplitude tremors.
That reflex factors may be important
relative to the displacement amplitude of
certain hand tremors is suggested by the
plots of Fig. 7A and B. These plots in-
dicate that, for rms displacements of the
tremor above about 100 pm, the rms de-
modulated EMG (calculated at the tremor
frequency) and the rms displacement of
the tremor are related by a power function.
The rms displacement amplitude increased
in proportion to the rms amplitude of the
EMG raised to approximately the 2.5 power
(an exponent between about 2.0 and 3.0).
It is of considerable interest that this
relation is very similar to that found (28)
for the soleus EMG and ankle tremor dis-
placement for a similar range of rms dis-
placements (a range of about three decades).
These plots for both ankle and hand tremor
(24, 28) indicate that the ratio of the EMG
to tremor displacement amplitude decreased
with increasing tremor displacement. It was
noted earlier (28) that, with increasing dis-
placement amplitude of cyclic muscle
stretch, a similar decrease occurred in the
sensitivity of primary endings of muscle
spindles (13, 18) of a decerebrate cat. The
finding that the EMG-tremor coherence de-
creases with frequency for small-displace-
 56 R. N. STILES
ment hand tremors (Fig. 6) also suggests
that rate-sensitive stretch receptors are
involved in the tremor-related neural
feedback.
While feedback from rate-sensitive muscle
spindles may be important in the mechanism
of certain hand tremors, it is not clear
whether this feedback normally functions
to stabilize the stretch reflex, as Merton
(14) suggested. The generally accepted role
of the stretch reflex in the mechanism of
small-amplitude tremors is that the feedback
sustains (through positive feedback) these
oscillations (9, 10, 12, 23). However, it
appears that this hypothesis does not con-
sider the possibility that these oscillations
may be maintained (the energy source) by
internal broad frequency band perturbations,
such as the asynchronous contractions of
active motor units. It now appears that the
system controlling hand position (or the sys-
tem controlling ankle rotation) cannot be
considered simply to be either stable (no
oscillations) or unstable (oscillatory). Rather,
this system seems to exist in many dif-
ferent states of stability. Also, it appears
from the results of several studies on fore-
arm tremor (3, 8, 11, 22) that this neuro-
muscular-load system also may exist in
many different states of stability, and that
these different states also depend on
combined mechanical and reflex factors.
A decrease in the EMG-to-tremor dis-
placement ratio with increasing displace-
ment amplitude of the tremor, coupled
with the assumption of internal disturbances
acting to maintain the tremor, suggests
that neural feedback may normally (for
small-amplitude tremors) act to attenuate
these oscillations. However, another im-
portant variable that should be considered
is the timing (or phase) between the motion
of the oscillating limb and the neural
feedback signal. (This quantity was cal-
culated but not reported in this study.) As
Rack (19) has noted, if the receptors are
sensitive to velocity (or acceleration) and
if the reflex delay is small, then the maxi-
mum reflex force may develop (depending
on the tremor period) while the muscle is
being lengthened. This reflex force would
oppose the motion and tend to either
abolish the tremor or reduce it in amplitude.
From the results of the present study (and
the similar study on ankle tremor (28)),
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it appears that these oscillations are never
abolished (as long as the muscles are con-
tracted). Rather, different steady-state con-
ditions apparently exist in which the internal
(and ex ternal) perturbation s are equally OP-
pose d by the combined mechanical and
reflex damping. However, these conditions
may be difficult to characterize fully since
any damping effect of a delayed neural
feedback signal will depend on both the
delay time and the period of the oscillation
(as well as upon the magnitude of the
resulting force). While the hand tremor
period was found to vary with increasing
displacement amplitude, the relation be-
tween these two quantities was not the
same for all subjec ts, nor for all m.ass loads
(see Fig. 4). Certainly the mean tension
of the muscles controlling the hand was not
controlled in the present study. As found
previously (27), the frequency of limb tremor
varies somewhat with this quantity. The
data of Fig. 8 indicate that varying the
mean position of the hand (reflecting a
change of the mean extensor muscle tension)
affected the tremor period in a way that is
consiste nt with a variable spring-mass
system.
In considering combined mechanical and
reflex factors in the mechanism of tremor,
the studies of Rack (19) and Stein (23) are
of interest. Both workers have sugge ,sted
models containing separate mechanical and
reflex oscillators, as well as a combined
mechanical-reflex oscillator mechanism, for
limb tremor. Each suggests that the stretch
reflex and the spring-mass systems normally
have separate natural frequencies of oscilla-
tion. Howe ver, under certain conditions
(mechanical load reflex gain, other factors),
these two systems may reinforce each
other such that a single-frequency, large-
amplitude oscillation develops, i.e., a
“limited instability” (11). While certain
features of this model may apply to the
data obtained in the present study, separate
mechanical and reflex oscillations were not
identified. Further, it is not clear whether
the combined mechanical-reflex oscillator
model presented by these workers (11, 23)
predicts the wide range of frequency and
displacement-amplitude values found for
both hand (Fig. 4; also see Ref. 24) and
ankle (28) tremor.
Auto- and cross-spectral analysis was
 MECHANICAL-REFLEX
performed with the assumption that the
grouping of electrical potentials of the
EDC and ECR muscles at the major tremor
frequency resulted from neural feedback
within the neuromuscular system control-
ling the hand. However, additional bands
with peak frequencies between about 9 and
15 Hz were occasionally found in the
EMG and acceleration spectra (see Fig. 2).
Elble and Randall (6) recently suggested
a central oscillator mechanism for the 9- to
lo-Hz hand oscillations that appeared when
the frequency of the major oscillation was
reduced with added mass (either 270 or 570 g
suspended from the hand). Their major
evidence was the failure of added mass
to reduce the frequency of this 9- to lo-Hz
oscillation and the presence of high co-
herence between the demodulated EDC
EMG and hand motion at this frequency.
In considering the mechanism of these
minor bands of the acceleration spectrum,
two points should be considered: 1) The
finding of high coherence between these
hand oscillations and the demodulated
EMG is not sufficient for support of a
central (CNS) source for the oscillator. 2)
The muscle-hand system is a distributed
mechanical system having more than one
mode of vibration because of the attach-
ment of more than one muscle. An addi-
tional factor of possible importance is
that oscillations of more proximal parts
(e.g., the muscle-forearm system) may be
transmitted by the hand (just as hand
oscillations may be transmitted by the
finger (6, 27)). The amplitude of any such
oscillation, as well as its transmission to
the hand, might be enhanced by increasing
the gravitational mass of the hand (gen-
erally resulting in an increased muscle
tension). Therefore, a mechanical-reflex
oscillator mechanism cannot be ruled out
as the source of these minor bands with
peak frequencies between 9 and 15 Hz that
are found in certain hand tremor spectra
(as well as in the wrist extensor EMG
spectra).
Certainly these minor tremor bands might
result from synchronized contractions of
single motor units that either fire at the
tremor rate (16, 17), or at rates higher
(5) or lower (16, 17) than that of the
tremor. The finding illustrated in Fig. 2A
and C, that these bands sometimes ap-
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OSCILLATOR HYPOTHESIS 57
peared in the EMG spectra of more than
one muscle controlling the oscillating limb,
suggests that motor units of different
muscles may have been synchronized in
some manner at these frequencies.
It is important to emphasize that the
presence of minor bands in the tremor
spectra seemed to depend on the subject
(they were found in only a few spectra
for certain of the nine subjects, and not
at all for two) and also on the displace-
ment amplitude of the major tremor band.
As the displacement amplitude of the major
band increased, these minor bands (if
present) generally disappeared from both
the acceleration and the EMG spectra
(also see Figs. 5 and 7 of Ref. 32).
While the mechanism of these 9- to 15-Hz
oscillations of hand acceleration is un-
certain, it seems that their presence had
only a minor effect (if any) on the dif-
ferent relations reported here for the
major band of the hand acceleration spectra.
The method used to induce hand tremors
of normal subjects at many different rms
displacement amplitudes above control
levels (above about lo-80 pm) was very
similar to that used previously to induce
ankle tremors at many different displace-
ment amplitudes (28). This method in-
volves the continuous maintenance by the
subject of the limb at a relatively con-
stant mean position against an opposing
gravitational force. While other methods
have been used to induce large-amplitude
oscillations of the forearm (3, 11) and
ankle (1, 28), these methods generally
resulted in a rather abrupt increase in the
tremor amplitude from near control levels
to very large amplitude levels (a “sus-
tained” tremor, or clonus). A general
problem with these methods is that the
intermediate-amplitude levels of the tremor
may not be recorded for analysis. There-
fore, the results obtained by these methods
tend to suggest that the neuromuscular
system controlling the particular limb has
only a few (perhaps one or two) states
in which oscillations occur (23).
In a previous study on hand tremor
from normal subjects, the frequency of the
tremor generally decreased as a power func-
tion with increasing displacement amplitude
for rms displacements between about 30-
30,000 ,um (24). In the present study, the
 R. N. STILES

tremor period (instead of log(frequency)) hypothesis for the mechanism of these

was plotted against the log(displacement)
amplitude of the tremor. These plots (Fig.
4 and others not shown) indicate that the
tremor period generally increased linearly
with log(displacement) for at least part of
the range of displacement amplitudes. (This
largely verifies the previous finding (24)
that showed a negative relation between
log(frequency) and log(displacement) over
most of the three-decade range of dis-
placement amplitudes.) However, outside
of these displacement ranges, the variation
of the period with log(displacement) ap-
peared to depend greatly on the subject
and the magnitude of the mass load. It
seems that, for tremors with rms displace-
ments above about 100 pm, no general
conclusion can be reached about the effect
of mass upon the tremor period and log(dis-
placement) relation. For certain subjects
and displacement amplitude levels (see Fig.
4B for subject 5), the tremor period
appears to be relatively independent of
mass loading.
During certain experiments, it appeared
that as the tremor amplitude increased
above certain levels (for certain subjects
such as subject 5) the mean contraction
level of the extensor muscles increased.
This was indicated by the tendency of
the mean position of the subject’s hand to
be maintained 5- 10” above the horizontal
as these large-displacement tremors oc-
curred. An increased mean tension of
the muscles resulting in an increased
muscle stiffness could explain the de-
creased (or relatively constant) tremor
period (see Fig. 8) that occurred with
increased displacement amplitude for cer-
tain subjects.
The difficulty in controlling mean tension
for large-displacement tremors also makes
it difficult to interpret the effects of mass
loading on hand tremors detected from
Parkinsonian subjects. As reported recently
(29, mass loading of the extended hand
of each of three Parkinsonian subjects
oscillations.
The very small effect (if any) of peripheral
loading on the frequency of the relatively
large-amplitude tremors for certain sub-
jects may help explain the failure of oscilla-
tions of pathological subjects to be en-
trained by a rhythmical force applied to
the limb (30, 31). It may be that neural
feedback factors may largely determine
the oscillatory behavior of the limb some-
what independent of peripheral loading or
forcing during the occurrence of large-
displacement amplitude tremors.
The results of the present study support
earlier findings (24, 28) that tremor oscilla-
tions of a limb of a normal subject can
occur at each of many different frequencies
and rms displacements. Certainly a system
producing these oscillations cannot be
viewed as existing in a relatively few
states of stability (or instability). Correla-
tion (coherence) between the demodulated
EMG and tremor was generally high for
all but the smallest amplitude oscillations.
However, this correlation appeared to
depend on the frequency as well as the
displacement amplitude of these tremors. In
general, the amplitude of the demodulated
EMG (calculated at the tremor frequency)
increased as a power function of the
increasing displacement amplitude of the
tremor. While the tremor period appeared
to be independent of mass loading for
certain subjects and displacement ampli-
tudes, the decrease in the period of
large-displacement tremors with increasing
extension of the wrist suggests that muscle
stiffness may be an important, uncontrolled,
factor in these experiments. Overall, the
results suggest that many different equi-
librium conditions can exist in which internal
and external perturbations, muscle-load
mechanics, and neural feedback effects
determine the major tremor oscillation of
the hand.
ACKNOWLEDGMENT

resulted in only a slight shift (if any) This study was supported by National Institutes
in the log(frequency) and log(displace- of Health Research Grant NS-14730.
ment) relation. Certainly a lack of an effect
of mass loading on the period of certain
large-displacement amplitude tremors (3 1) Received 27 August 1979; accepted in final form 21
cannot be used to support a central oscillator January 1980.

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 MECHANICAL-REFLEX OSCILLATOR HYPOTHESIS 59

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