NEGATIVE ADAPTATION AND REFRACTORY

PHASE IN THE EYELID REFLEX »•2

BY LOUIS H. COHEN
Yale University

The diminution and eventual disappearance of a reflex

response consequent to repeated stimulation (negative adap-
tation) has been studied by Griffith (7), Dodge (4) (5), Cason
(2), Dodge and Louttit (6) and the writer (3). Griffith (7)
and Dodge (4) have shown that this phenomenon exists in
post-rotation nystagmus and compensatory eye-movements,
and Dodge has demonstrated a "typical" curve which drops
rapidly at first with subsequent negative acceleration. Nega-
tive adaptation of the eyelid reflex has been demonstrated by
Cason (2), and by Dodge (5). In the knee-jerk, however,
Dodge has shown that diminished amplitude of response to
repeated stimulation is extremely small over a period of some
20 months.
Dodge and Louttit (6), in investigating the body-start
reflex and the ear reflex of the guinea pig to sudden noise,
demonstrated that the speed of negative adaptation to re-
peated stimulation was greater for the former reflex than for
the latter. They attributed this difference to the fact that
the relative refractory phase of the body-start reflex is longer
than that of the ear reflex. With this experiment as a basis,
the writer (3) studied variation in the rate of negative adap-
tation of the body-start reflex of the guinea pig when stimu-
lation occurred at different intervals in the refractory phase.
It was shown in that investigation that there is a relationship
between the speed of negative adaptation and various inter-
vals within the refractory phase of the reflex at which stimu-
lation falls, i.e., there were points of greater and smaller
1
Grateful appreciation is due to Professor Raymond Dodge and Professor L. T.
Spencer for frequent advice and criticism during the course of this investigation.
* This paper is part of a dissertation submitted to the Faculty of the Graduate
School of Yale University in candidacy for the degree of Doctor of Philosophy, 1929.
447
448 LOUIS H. COHEN

susceptibility to negative adaptation which followed the form

of a modal curve.
The present investigation is an extension of the inquiry
into the relationship between negative adaptation and exci-
tability of response at various moments in the refractory
phase. The eyelid reflex was studied because information
regarding the recovery of excitability during its refractory
phase was at hand (2), and because of available recording
techniques.
APPARATUS

The apparatus consisted essentially of a camera which recorded the movements

of a light beam, reflected from a mirror on a lever attached to the eyelid. Noise stimuli
were provided by wire hammers released electromagnetically at appropriate moments
against a sounding box.3
PROCEDURE AND RESULTS

Fifty-five adult subjects, of whom 26 were women, were apportioned by chance

to II stimulus-interval groups of 5 subjects each. The stimulus-intervals selected
were o, 20 sigma, 40 sigma, 80 sigma, 160 sigma, 320 sigma, 640 sigma, 1 second, 2
seconds, 3 seconds, and 4 seconds. Sixty trials, spaced at 30 second intervals, were
given for each subject. The subjects were unaware of the number of trials to be re-
corded. Except for the slight sounds made in adjusting the apparatus, and occasional
introspective reports, the laboratory was relatively silent, except, of course, at mo-
ments of stimulation.

Table 1 presents average amplitudes of Ri and R2, re-

sponses to the two stimuli at the given interval, of five con-
secutive trials for each group throughout the series.
(1) It may be seen that data of R2 were not available for
purposes of statistical treatment until the 320 sigma interval.
Although occasional responses to the second stimulus were
evoked at the 40 sigma, 80 sigma, and 160 sigma stimulus-
intervals, they were of such small amplitude that only a quali-
tative record could be made of their existence. For example,
at 40 sigma, R2 occurred only three times in the 300 trials
recorded. The number of i?2's was greater at 80 sigma, and
still greater at 160 sigma.
(2) Both Ri and R2 for each group undergo a progressive
negative adaptation. This confirms previous findings as to
the effect of repetition of stimulation on reflex response.
(3) Negative adaptation of Ri at the 40 sigma and 80
•For a more extensive description of the apparatus the reader is referred to (2).
 NEGATIVE ADAPTATION IN EYELID REFLEX 449
TABLE i
AVERAGE AMPLITUDES (IN MM.) OF RI AND RI FOR ALL STIMULUS-INTERVALS
(60 TRIALS)

Trial Zero 20<r 40<r 8o» I6O<T 320V 6407 1 tec. 2aec 3 set 4aec

i-s 14.I 18.8 17 .8

19.1 2O.4 11.4 i?.o 12.8 11.7 13-9 11.7
6-10 9-7 14.7 17.8 I2.O
18.8 8.6 4-4 9.1 6.9
u-iS
16-20
9.0 IO.8 19- <; 174 12.7
84 1 0 . <; 13.4 16.5 10.1
li
6.0
9-9
8.8
7-7
6.0
.8
.6
0-4
7A
9-8
10.4
21-25 .5-6 9-9 I.S-2 IS-3 10.6 2.8 9-1 4-5 .2 6.9 9.8
p. 26-30 5.0 9.1 10.2 17.1 9-3 4.4 9-9 S-9 •3 3.6 9-7
31-35 4.6 »-5 12.3 14.0 9-9 2.7 7-1 6.5 .1 3-2 7.6
36-40 4-4 7-5 12.2 17.1 9.1 2.4 8.1 4.2 .0 2.2 8.4
4I-4S 5-4 7.8 10.2 17.2 9-5 2.7 8-3 S-4 •3 3-9 11.7
46-50 4-3 7-7 133 16.0 8.8 1.9 5.8 4.8 .0 4.0 12.0
4.4 6.9 10.2 12.5 9.4 6-3 5-1 .0 1-7 10.8
si-ss
56-60 4.1 6.8 11.1 13-4 8.4
1-3
1-3 4.8 4-7 .0 2 . 1 10.1

1.8 84 3.6 3.0 10.7 114

6-1 o 6.1 2.4 •4 9-3
-5 3-9 2.5 .2 4^6 9-3
16-20 4-9 .0 5-2 10.2
21-25 5-8 1.6 .0 1-9 9-4
26-30 .6 3-7 24 .1 •7 10.0
31-35 •3 3-4 2.8 .0 1-4
36-40 •9 2.7 .0 •4 8.8
41-45 •7 4.6 2.8 .2 •4 8.7
46-50 .1 3.2 2-5 .0 -7 10.2
.2 2.0 2.9 .0 104
56-60 .2 2.7
.0
i 9.6
8.9

sigma stimulus-intervals is much more irregular than that at

other stimulus-intervals.
(4) There is a differential amplitude decrement depend-
ent, apparently, on stimulus-interval. Table 2 presents the
TABLE 2
TOTAL PERCENTAGE OF NEGATIVE ADAPTATION OF RI AND RI FOR ALL STIMULUS-
INTERVALS (PERCENTAGE DIFFERENCE BETWEEN AMPLITUDE OF
FIRST AND LAST FIVE TRIALS)

Stimulus-Interval Ri Rt
zero 70.9
20 sigma 62.7
40 sigma 37.7
80 sigma 31.3
160 sigma 58.8
320 sigma 88.6 88.9
640 sigma 68.0 82.2
1 second 63.3 25.0
2 seconds 100.0 100.0
3 seconds 84.8 944
4 seconds 13.7 22.0
4SO LOUIS H. COHEN

percentage differences between the average of the first five

trials and the last five trials for each response.
The relative speed of negative adaptation for each stimu-
lus-interval may, therefore, be summarized as follows: the
greatest adaptation is undergone by the group at the stimulus-
interval of 2 seconds, followed in order of lessening adaptation
by the groups at 3 seconds, 320 sigma, zero, 640 sigma, I
second, 20 sigma, 160 sigma, 40 sigma, 80 sigma, 4 seconds.
Adaptation of i?2 is also greatest at 2 seconds, followed in
order of decreasing adaptation by the groups at 3 seconds,
320 sigma, 640 sigma, 1 second, 4 seconds, demonstrating
absolute agreement with differential adaptation of Rx. R2's
for the zero, 20 sigma, 40 sigma, 80 sigma, and 160 sigma
stimulus-intervals are not presented because of lack of suffi-
cient quantitative data.
Order seems to be brought out of the apparent chaos of
the alignment of these series if the curve of differential adap-
tation (Fig. 1) is examined. This graph demonstrates the
relationship between stimulus-interval and percentage of adap-
tation as given in Table 2. In this curve the following fea-
tures of significance are suggested:
(1) Effectiveness of stimuli in producing negative adap-
tation is greater as the interval between stimuli is increased
from zero to 80 sigma.
(2) The curve of negative adaptation follows a definite
wave-like form with low points (resistance to adaptation) at
80 sigma, 640 sigma, 1 second, and 4 seconds; and high points
(susceptibility to adaptation) at 320 sigma and 2 seconds.
Intervening points make this curve a fairly smooth one: it
will be noted that the periods of successive phases are increas-
ingly greater.
(3) Negative adaptation of R2 follows a wave-like curve
fairly homologous with that of the Rx curve.4
1
It is interesting to note that the correlation coefficient (p) between negative
adaptation of Jit and At is +.943; when converted to the product moment r it is +.947,
with a P.E. of ±.029. Of course, there are only 6 cases represented in these figures,
but the close relationship is suggested nevertheless. This phenomenon, together with
other data of a similar nature, will be discussed in a later paper.
 NEGATIVE ADAPTATION IN EYELID REFLEX

DISCUSSION

In a previous paper (2) the recovery of excitability during

the refractory phase of the eyelid reflex was presented. I t
was seen that this curve was wave-like in form, i.e., that the
recovery of excitability was not in simple arithmetical pro-
gression, but rather was a complex rhythm of alternating
recovery and depression.
The comparison of the wave-like curves of recovery of
excitability of the three subjects in that investigation and an
apparently homologous curve of adaptation of the fifty-five
subjects of this investigation seems to be demanded. If the
tables presenting the data of that investigation are examined,
therefore, it will be seen that the mean average percentage
amplitudes of R2 (calculated from the R2/R1 ratio) are related
to the total percentage of negative adaptation of response for
the subjects of the present investigation in an inverse manner.
The relevant portions of these tables are presented again for
convenience.

Total Percentage of
Negative Adaptation of R Average Percentage of
for Stimulus-Intervals Ri for 3 Subjects for
Stimulus-Interval of 320 sig.-4 sec. Stimulus-Intervals
of 320 sig.-4 s e c
Rt &

320 sigma 88.6 88.9 33-8

640 sigma 68.0 82.2 55-1
I second 63-3 2J.O 72.7
2 seconds 100. IOO. 50.0
3 seconds 84.8 94.4 62.9
4 seconds 13-7 2Z.O 86.2

It may be seen that although the direction of this inverse

relationship between negative adaptation and excitability is
consistent for these stimulus-intervals, absolute amounts of
concomitant variation do not follow the above generalization.
Thus, assuming the rough quantitative proportionality of the
negative adaptation-excitability ratios at 320 sigma, 640
sigma, and 1 second to be correct, one would expect at the 2
second stimulus-interval either a smaller percentage of nega-
tive adaptation for the obtained percentage of excitability, or
452 LOUIS H. COHEN

the obverse, i.e., a smaller percentage of negative adaptation.

This same difficulty exists at the 3 second stimulus-interval.
The 4 second stimulus-interval, however, preserves the quan-
titative aspect of the generalization. The differential nega-
tive adaptation of response seems to follow, therefore, a curve
of wave-like form of a nature reciprocal with the homologous
g
s too
The Relationship between Hegatire Adaptation ana Sttaulud-

Stimulus-Interval
•r HO. IK. XMC.
II j U ^
br Fig- X

curve of excitability. The correspondence of these two curves

argues for the validity of the statement that one may be a
function of the other.5
It was seen that the rate of negative adaptation was found
to decrease as the interval between stimuli from zero to 80
sigma was increased. If the reciprocal relationship of exci-
tability of R2 and negative adaptation is true, these data
would indicate a progressive increase of excitability as the
intervals between stimuli are increased up to 80 sigma, despite
the absence of recorded secondary response.
Why should this relationship between excitability of the
reaction-system and its speed of negative adaptation be in-
verse? Our data demonstrate this, and there seem to be
theoretical reasons also for its validity, namely, the refractory
phase theory of inhibition and, by analogy, Heymans' Law
of Inhibition.
Briefly, the refractory phase theory of inhibition states
'The correlation coefficient (p) between negative adaptation of .Ri and excita-
bility was found to be —.89 (f = —.90, P.E. = ±.053) and that "between negative
adaptation of Si and excitability to be —.71 (r = —-73, P-E. = ±.134). Since n
for both these figures is only 6, the values may have limited significance, but the
mathematical relationship of the variables is, nevertheless, demonstrable.
 NEGATIVE ADAPTATION IN EYELID REFLEX ' 453

that inhibition is set up by the falling of successive stimuli

within the refractory phase of the stimuli which precede them.
If the stimuli are given at a fast rate, total inhibition is
achieved because of the absolute refractory phase; if the stim-
uli are presented at a slower rate, involving the relative re-
fractory phase, various degrees of partial inhibition may be
set up; if the stimuli are so presented that they fall within the
rebound of previous stimuli, the effect is not inhibition, but
summation. If one may consider negative adaptation as a
type of protracted inhibition, it follows that stimuli spaced at
intervals of low excitability will undergo greatest negative
adaptation (inhibition) which will be less complete the greater
the excitability of the reaction-system. Furthermore, stimu-
lation in the supernormal phase, by nature of the summation
of excitability set up, would be most resistant to modification
in this form. Obviously, the relevance of these experiments
to the refractory phase theory can be assured only by verify-
ing the two assumptions implied; viz., that excitability as used
in the present sense is truly a case of excitability or irritability
as physiologically understood; and secondly, that negative
adaptation is a case of protracted inhibition. If the above
reasoning and its assumptions are correct, the present investi-
gation points to the significance of the refractory phase in
producing inhibition.
This inverse relationship between excitability and negative
adaptation (inhibition) is not without precedent in the psy-
chological literature. In investigating the validity of Hey-
mans' Law, Spencer (8) found a correlation of +.907 between
visual threshold and its susceptibility to inhibition for 50 sub-
jects, and Spencer and Cohen (9) demonstrated a correlation
of +.955 and +.968 for these same variables for 50 cases in a
single subject, these coefficients running as high as +.992 in
the last half of the experiment. In other words, both for the
group and for the individual, thresholds were higher as sus-
ceptibility to inhibition was greater, increased thresholds, of
course, meaning decreased excitability. Here the assump-
tions must be made that excitability of a reflex reaction-system
at various moments within the relative refractory phase is
30
454 LOUIS H. COHEN

analogous to sensory threshold, and that negative adaptation

is analogous to sensory inhibition. Since excitability (thres-
hold) was found to be correlated to an almost perfect degree
with susceptibility to inhibition in accordance with Heymans'
Law, an analogous phenomenon in the present data would
seem to fall within that generalization.
CONCLUSION
From these data it seems evident that the speed of nega-
tive adaptation of reflex response is a function of the degree
of excitability of the reflex at given moments in the refractory
phase. These two variables were found to be reciprocal, thus
indicating a mathematical relationship between speed of ex-
tinction of reflex response and specific excitability of the reflex
reaction-system. The validity of this inverse relationship is
argued ex post facto from the refractory phase theory of
inhibition and Heymans' Law.
REFERENCES
1. CASON, H., / . Exper. Psyckol., 5, 1922, 153-196.
2. COHEN, L. H., / . Exper. PsychoL, 15, 1932, 436-446.
3. , / . Comp. PsychoL, 9, 1929, 1-16.
4. DODGE, R., / . Exper. PsychoL, 6, 1923, 1-35.
5. , Elementary Conditions of Human Variability, Columbia University Press,
New York, 1927.
6. AND LOUTTTT, C. M., / . Comp. PsychoL, 6, 1926, 267-285.
7. GRIFFITH, C. R., / . Exper. PsychoL, 3, 1920, 89-125.
8. SPENCER, L. T., / . Exper. PsychoL, 11, 1928, 88-97.
9. AND COHEN, L. H., / . Exper. PsychoL, 11, 1928, 194-201.

(Manuscript received June 23, 1931)