THE RELATIONSHIP BETWEEN REFRACTORY PHASE

AND NEGATIVE ADAPTATION IN

REFLEX RESPONSE I»
LOUIS H. COHEN
Yak University
The factors determining the diminution and eventual dis-
appearance of a reflex response consequent to repeated stimula-
tion have received little experimental consideration. Griffith2
and Dodge* have shown that the amplitude of post-rotation
nystagmus undergoes a rapid diurnal decrease with repeated
stimulation, compensatory eye movements displaying a similar
adaptation to rotation. The effect of repetition of stimuli for
these two factors of vestibular reflex response was therefore to
induce a modification in the form of negative adaptation.
Griffith demonstrated that such experimental conditions as time
of day, amount and number of rest intervals between stimulations
and between series, general organic condition of the subject,
and "transfer" were significant in determining the speed of
reduction of response. Little is known concerning negative
adaptation of reflex response when the reaction system is in a
state of diminished excitability or relative refractory phase.
That there might be some relationship between this response
modification and the relative excitability of the given reaction
system seemed plausible in the light of the recent work of Dodge
and Louttit* who recorded the body start reflex and the ear
reflex of the guinea pig to sudden noise.
1
Both for the problem and the general experimental' method the writer is
indebted to Professor Raymond Dodge. The writer takes much pleasure in
expressing his gratitude to him and to Professor L. T. Spencer for their frequent
advice and invaluable criticism.
1
Griffith, C. R.: Jour. Exp. Psychol., iii, 1,1920.
»Dodge, R.: Jour. Exp. Psychol., vi, 1,1923.
4
Dodge, R., and Louttit, C. M.: Jour. Comp. Psychol., vi, 3,1926.
1

COUPABATITB FSTCHOLOGT, VOL. IXi HO. 1

 2 LOXTIS H. COHEN

This body start reflex consists of a slight thrust of the forelegs

downward, followed by a retraction of the forelegs. Dodge and
Louttit state that "the length of the refractory phase for the
body (start) reflex is shown by the records to be over 0.5 second.
Direct observation showed that it may last 2 to 3 seconds . . . .
though this is a tentative estimate based on unrecorded observa-
tions" (p. 282). The latency of reaction, though subject to
individual differences, was shown to last about 30 sigma. A
photographic method of registration was employed.
Dodge and Louttit demonstrated further that on 3 guinea
pigs, with 30 successive trials, each consisting of 2 stimuli ap-
proximately 0.5 second apart, the amplitude of response of this
start reflex tends to become increasingly smaller. They showed
that the start reflex, whose relative refractory phase was longer
than that of the ear reflex, tended to become eliminated sooner
from the reflex pattern than the latter of shorter relative refrac-
tory phase, although the amplitude of both became less with
repeated stimulation. Differential rate of negative adaptation
seemed to be associated with the length of the relative refractory
phase of the two respective reflexes studied.
If the differential rate of negative adaptation is associated
with the length of the relative refractory phase when a series
of stimuli are given at a certain rate, it is conceivable that it is
because the second and succeeding stimuli, for responses of
relatively short refractory phase, occur late in the phase, while
for responses of longer refractory phase the second stimulus occurs
relatively early (i.e. nearer the absolute refractory phase). If
this is so, it may be that with a given reflex a variation in the
rate of adaptation will be found when stimulation occurs at
different intervals in the refractory phase.

The present problem is concerned chiefly with the effect of

repeated stimulation on the differential rate and amount of
negative adaptation of the body-reflex of the guinea pig as
dependent on the intervals within the relative refractory phase
at which the second stimulus occurs.
 a

FIG. 1
 LOUIS H. COHEN

APPARATUS
The apparatus for the present experiment involves a means of
graphically recording the guinea pigs' body start reifex to two
temporally controllable sound stimuli. Silence of operation other
than at moments of stimulation was therefore necessary.
Sound stimuli were produced by the successive striking of two
spring-driven steel mouse-traps against the closed side of a
sounding box after an excursion of 90 degrees. This device
has been used by Dodge in his pendulum photochronograph.*
Each of the mouse-traps was "held down" by the cores of sole-
noids when the circuit through each solenoid was closed. It
may be seen, therefore, that the breaking of each circuit at ap-
propriate moments would release each of these electromagnetic
"triggers," which on drawing back would release the mouse-trap
springs. The moment of impact was recorded on a kymograph
by a pointer on a long wooden lever attached to the sounding
box. The movements translated to this recording lever were
caused by the slight movements of the sounding box when struck
by the spring traps. The struck side of the box was reinforced
by a piece of tin; the intensity and quality of both noises ap-
peared to be equal.
Connected in series in the circuit to each solenoid was a
contact consisting of a short brass rod with one lead, resting on an
upright brass strip forming the other lead, the circuit being
closed when the contact points met. The slightest upward
movement of the brass rod would remove it from the strip, thus
breaking the circuit. Each contact was supported by a 5 inch
lever arm which was attached at its other end to a common
center. One arm was stationary, the other movable throughout
the circumference of a circle 10 inches in diameter. Thus, the
movable arm, and therefore its corresponding contact, could
be adjusted to any arcuate distance relative to the first. A
360 degree protractor facilitated the reading of distances in
degrees between contacts.
The breaking of the circuits was accomplished by placing an
6
Dodge, R.: Jour. Exp. Psychol., ix, 2, 1926.
 REFRACTOKY PHASE AND NEGATIVE ADAPTATION 0

offset on a rotating kymograph, which during the course of ite

revolution would strike the upright brass rod of each contact in
turn. Sound of the impact of this offset on the brass rods was
minimized to inaudibility by enclosing the brass rods in rubber
tubing except, of course, at the points of electrical contact.
The kymograph was connected by a belt to an approximately
silent spring phonograph motor.
That this motor possessed the further virtue of constant speed
once its maximum momentum was reached (about 0.25 second
after starting) was shown by the fact that the desired time in-
tervals between stimuli increased in the same arithmetic progres-
sion as the degree distances between contacts, with a mean
variation of not more than 1 per cent, as measured in hundredths
of a second. Thus, a 1- second interval between stimuli was pro-
cured empirically by placing the indicators of the contacts 70
degrees apart at a certain speed of the motor. By placing the
contacts 140 degrees apart, a 2-second interval was procured;
210 degrees apart, a 3-second interval; 280 degrees apart, a 4-
second interval. This empirical method was checked by the
mathematical determination of the speed of the distal end of the
offset as determined by its radius and the speed of the motor.
The revolution of the motor automatically turned the kymograph
and the offset, the time interval between stimuli being therefore
as accurate as the speed of the motor.
The body movement recorder was practically the same as that
used by Dodge and Louttit, the same basket in fact being used
with some slight modification.6 This basket was 24 by 13 by 7.5
cm., made from sheet aluminum a little less than 1 mm. thick,
and weighing 325 grams. A friction surface was afforded the
feet of the animal by a thin sheet cork floor. This had short
upright wooden cleats which were so placed that the apparent
center of gravity of the guinea pig would come directly above the
bearing supporting the basket from underneath. The ends of
this bearing were fitted into point supports, movements of the
basket therefore being in a plane perpendicular to its horizontal
• Op. cit., p. 269.
6 LOTTIS H. COHEN

axis. The basket was held in elastic equilibrium by two adjust-

able rubber springs from above, one at each end. The damping
effect of the springs was increased by a loose fitting oil plunger
below the basket. If any movement greater than the inertia of
the basket, plunger, and springs disturbed this equilibrium, the
extent and duration of the disturbance was recorded on the
kymograph by a recording pointer of wood 17 cm. long, projecting
from the top of the basket, forming a continuation of its upper and
inner edge. The distance from the bearings of the basket to its
recording segment was 28 cm. The sensitivity of the balanced
basket was determined in the manner employed by Dodge and
Louttit; 7 10-, 20- and 50-gram weights were placed on the front
end of the basket, and a kymograph record taken of the displace-
ment of the pointer upward when the weights were quickly
TABLE 1
Calibration of the balance recorder
WEIGHTS DISPLACEMENT OF POINTEB

grama mm.
10 12
20 24
50 60

removed. This calibration of the basket shows not only the

sensitivity of the balance, but also a constant relationship
between gram weight and millimeter displacement of the pointer.
The perpendicular distance from the sounding box to the front
cleat of the basket against which the forefeet of the pig rested
was 64 cm.; from the sounding board to a point between the ears
of the guinea pig was a distance of about 50 cm.
A fairly accurate method for time recording was necessary for
the purpose of empirically checking the time interval between
stimuli and for measuring simultaneously rate and duration of
body movement. In order that the requirement of silence be
fulfilled, the timing method employed consisted in connecting
two 100 d.v. electrically driven tuning forks in series, the record-
ing fork having its contact breaker removed. The interrupting
7
Op. cit., p. 270.
 EEPRACTOBY PHASE AND NEGATIVE ADAPTATION 7

fork was placed outside the laboratory. No sound from the

recording fork could be heard except when it was placed very
close to the ear.8 At the completion of a given trial, the drum was
removed from contact with the markers by turning a spiral gear.
It could then be raised without interference from the markers, so
that the succeeding trial could be recorded directly beneath its
preceding trial when the drum was brought back to the markers.
The relative position of the markers thus remained unchanged.
Eight male guinea pigs were used as subjects.8
PROCEDURE
The eight guinea pigs were divided into 4 groups of 2 each.
Classification into groups was made on the basis of weight, so
that the mean weight of the groups might be approximately the
same.
Each of these pigs was experimentally subjected to identical
conditions except that for Group A the interval between stimuli
of a stimulus pair was 1 second; for Group B, 2 seconds; for
Group C, 3 seconds; for Group D, 4 seconds. There seemed to be
prima facie evidence that the 4- second interval was still in
refractory phase as shown by the diminished second response by
both pigs of Group D as recorded in the first 10 trials. (See
table 2.) Thus, the second stimulus was timed so that in all
cases it fell apparently within the relative refractory phase, closest
to the absolute refractory phase for Group A and farthest from
the absolute refractory phase for Group D. The first stimulus
in every case was administered after the kymograph had reached
its maximum momentum.
Each pig was stimulated 10 times per day for twenty con-
secutive days,10 a 30-second interval being allowed to elapse
between the moment of the second stimulus and the beginning
of the succeeding trial. The experimentation occurred at
approximately the same time each day.
* This timing method was suggested by Professor Spencer.
• These reactors were made available through the courtesy of the Committee
for Research on Sex Problems, of the National Research Council.
10
In the case of Group A, this procedure was extended to 30 days for a total of
300 trials. This will be discussed later.
8 LOTJIS H. COHEN

TABLE 2
First three amplitudes of displacements for each subject {in miUimeters)
Ai At Bi B> a Ct Di D:

Ri Downthrust

1 0.2 0.9 0.6 0.6 0.5 0.2 0.1 0.0

2 0.2 0.9 0.7 0.4 0.3 2.0 0.2 0.2
3 0.3 0.8 0.8 1.2 0.0 0.3 2.0 0.0
Mean 0.23 0.87 0.70 0.73 0.27 0.83 0.77 0.07
Group
mean... 0.55 0.72 0.55 0.42
Retraction

CO CO CO
1 1.0 4.0 1.1 0.2 0.2 0.2 0.3

o o o
2 0.2 3.0 1.1 0.4 0.4 0.1 1.0
3 0.4 0.3 1.0 0.3 3.0 0.5 1.0
Mean 0.53 2.43 1.07 0.3 1.2 0.27 0.3 0.77
Group
mean... 1.48 0.69 0.74 0.54
Ra Downthrust

en to d
1 0.1 0.1 0.2 0.1 0.3 0.2 0.0

o o o
2 0.2 0.1 0.5 0.1 0.1 0.3 0.1
3 0.1 0.1 0.3 0.0 0.3 0.5 0.0
Mean 0.13 0.1 0.33 0.07 0.23 0.33 0.23 0.03
Group
mean... 0.12 0.20 0.28 0.13
Retraction
1 0.1 0.1 0.2 0.3 0.2 0.2 0.0 0.2
2 0.1 0.1 1.0 0.1 0.5 0.2 0.2 0.1
3 0.1 0.1 0.6 0.0 0.2 0.3 0.1 0.3
Mean 0.1 0.1 0.6 0.13 0.3. 0.23 0.1 0.2
Group
mean... 0.10 0.37 0.27 0.15
 EEFRACTOET PHASE AND NEGATIVE ADAPTATION 9

After the contacts were closed and the spring mouse traps
"set" by the magnetized solenoid triggers, the experimenter
started the motor, which in driving the kymograph broke the
circuits through each solenoid at appropriate moments, a con-
tinuous record of time in hundredths of a second, of the stimulus,
and of the body movement of the subject thus being made until
the motor was stopped.
The data procured from the record of the reactions of the
subjects under the conditions of the procedure indicated above,
include, for any one trial, two separate responses, the second of
which presumably falls within the relative refractory phase at a
point differing with the group of the particular subject. It will
be remembered that the body-response of the guinea pig to sudden
noise is a thrust of the forelegs downward, followed by a retrac-
tion of the forelegs. Doubtless there are other factors in this
reflex pattern. The present problem, however, concerns itself
merely with these two grosser and experimentally more accessible
components. For each response to the two sound stimuli, the
following readings were taken:
1. Latency of response in estimated time units of one sigma.
2. Amplitude of downthrust as read to the nearest tenth of a
millimeter.
3. Duration of downthrust in estimated time units of one sigma.
4. Amplitude of retraction from maximal displacement of
downthrust as read to the nearest tenth of a millimeter. When
preceded by the downthrust, there is apparently no separation
between downthrust and retraction, indicating that normally the
two are component phases of a single total reaction.
5. Duration of retraction in estimated time units of one sigma.
It must be emphasized that these data are not absolute values.
The apparatus conditions make relative data alone accurate.
How the apparatus modifies the rate and extent of displacement
amplitudes has been previously noted.
The data are not distributed evenly over the 20 days of the
experiment, since they are selected and often include less than the
actual 10 trials per day. This "selection" of data was made
necessary by the fact that if any movement was recorded before
 10 LOOTS H. COHEN

the moment of either stimulation, that particular response was

entirely disregarded, because the movement changed the position
of the pointer, not only from that of other trials, but also from
the position of the pointer in the other responses of the same trial.
These changes in the position of the pointer were made by shifts
in the center of gravity of the subject. This occasional movement
of a nature similar to the responses studied is interesting to note,
since it seems to demonstrate that closely allied responses can
occur while the response elicited by a prior stimulus is in relative
refractory phase. This accidental movement was not prevented
in order to maintain the greater desideratum of unhampered
movement.
RESULTS
It was found that the range of latencies of response is from the
order of 22 sigma to the order of 54 sigma, the central tendency
being at about 35 sigma. This value agrees fairly closely with the
latencies found by Dodge and Louttit. The latencies of the
second response were found to be appreciably greater than those
of the first response, which suggests the findings of Lucas,11
who showed that responses in relative refractory phase had longer
latencies the sooner the first stimulus was followed by the second.
No such relationship was worked out here because of the limi-
tations of these particular data, principally because of the wide
variation, but there seemed to be some differential factor operat-
ing here also which produced the prolonged retardation of
response occurring in relative refractory phase.
The initial amplitudes of movement of the first and second
responses (Ri) and (R2) are presented in table 2 which includes
only amplitudes of the first three trials and their respective means.
It may be seen that amplitude of retraction for each pig is in
general somewhat greater than amplitude of downthrust. Fur-
thermore, the group means for Ri are fairly similar, indicating
that the initial energy of contraction is on the average the same
for all groups. The duration of the respective amplitudes was
" Cit. by Verwora, M.: Irritability, New Haven, 1913, p. 160,161.
 REFRACTORY PHASE AND NEGATIVE ADAPTATION 11
found to vary as the amplitude. R* is less in amplitude than Ri;
the differences between the displacements of Rg and those of Ri
seem to be greatest for subjects Ax and A* and least for subjects
Ci and Cs. The amplitude of response is, in general, shown to be

Dtaptaoatunt upUtiadM of Rl far Groupi A, uplltudM of 82 fur Omaf k,

B, 0, Bid -Sublet** Di art 02
in Intamls of * Day* In i s U m d * of A Day*
femthnut R»tr»rtlcn ttanrthrurt "*"

.9

FIG. 2 Fia. 3

the less the sooner the second stimulus succeeded the first, which
corroborates the usual findings in relative refractory phase.
The data of displacement amplitudes of both subjects in a given
group (except group D) were combined in order that group
 12 LOUIS H. COHEN

comparisons might be made. For the sake of brevity, the data

are presented in curves rather than tables (figs. '2 and 3).
These curves have been smoothed by the averaging of five
contiguous values.
It may be seen that in general, amplitudes of Ri and Rs decrease
for each subject at different rates. Despite the similarity of the
magnitudes of the intitial amplitudes of response (table 2)
there is a differential rate of reduction of response within the
first four-day interval which persists throughout the 20 days of
experimentation. In group C there are greater decreases in
amplitude than in group B, which however are less than the
decreases displayed by group A. In the case of subject Di
the decreases are greater than those of any others. With the
exception of the amplitude of retraction of Ri, subject D2 shows
no decrease whatever in succeeding trials. It will be remembered
that subject D* originally displayed an R s apparently within the
relative refractory phase. Continued stimulation may therefore
have artificially shortened the relative refractory phase, so that
the second stimulus occurred in the rebound with the effect
noted. With this exception it seems, therefore, that the later the
second stimulus occurs in relative refractory phase, the more
effective are both the first and the second stimuli in producing
negative adaptation.
This differential rate and amount of negative adaptation will
be made clearer perhaps in considering the percentage frequency
of responses of whatever amplitude occurred on successive days,
viz: downthrust, retraction, and either. Graphs representing
these data (figs. 4 and 5) include in broken line the separated
data of subjects Di and D8 to make clear the dissimilarity between
these two subjects and the apparently different physiological
conditions under which subject D2 was operating.
It may be seen that the frequency of complete absence of re-
sponse in the first four days is not very great or distinctive for
most subjects, indicating that despite the large decrease in am-
plitude of movement as previously shown, complete absence of
movement is rarely obtained within this first interval. The
general relationships between groups A, B, C and subjects Di
 REFRACTORY PHASE AND NEGATIVE ADAPTATION 13

Percentage Frequency (Vertical Axes) of Amplitudes

of Ri and R2 for Groups A, B, C, end Subjects Di
and D2 in intervals of 4 Days.
R Downthrust
100 100
80 80
60 60
40 40
20
20

4 § 12 16 20 8 12 16 20
Days

Retraction
100
80

.60

40

20

6 12 16 20
Day.

Sitter R2
100 * k
-<:
80

60

40

20
\
V.
i k2L 10 2U
» a Days

FIG. 4
14 LOUIS H. COHEN

and D s previously shown are, however, corroborated by these

data.
That in the case of subject D s the second stimulus occurred
after the relative refractory phase was over and perhaps in the
hyper-excitable period of the rebound has been already suggested.
For subject Di the second stimulus occurred possibly on the
"edge" of relative refractory phase or perhaps during a period of

50
40

50

60

70

80

90

100

FIG. 5. RELATIONSHIP or SECONDS INTERVAL BETWEEN STIMULI (HORIZONTAL

AXIS) TO AVERAGE PERCENTAGE FREQUENCY OF NEGATIVE ADAPTATION
(VERTICAL AXIS) OF R* AND RJ FOR GROUPS A, B, C, AND
SUBJECTS D I AND D S

recovered excitability in the rebound. Individual differences in

the length of the relative refractory phase therefore seem to exist.
A curve demonstrating the relationship between effectiveness
of both stimuli in producing negative adaptation, and the interval
between stimuli is presented in figure 5. Although the points
at the 4- second interval represent the data of but one subject
and points at the other intervals of but two subjects, there is an
apparent modal curve between these factors which may be of some
 HEFBACTOKY PHASE AND NEGATIVE ADAPTATION 15

significance in the demonstration of susceptibility of reflex re-

sponse to adaptation and may suggest an optimal point for the
production of adaptation.
For group A, 100 more trials were run under the same experi-
mental conditions, but there was no change brought about in
these additional trials which suggested any tendency toward
further reduction of response.
The ear reflex in this experimental situation seemed to the
observer to be subject to a consistent decrease in amplitude of
movement for all reactors. In no case was complete absence of
movement noted. No objective record of the ear movements
was taken, but it seemed that complete negative adaptation of
this reflex would require a far greater number of trials than neces-
sary for the body reflex, to attain a specific level. No differential
speed in this decreasing amplitude of ear movement, of course,
could be noted among any of the reactors, since these observations
were merely casual.
STJMMABY AND CONCLUSIONS

Eight male guinea pigs were separated into 4 groups of 2 each

and stimulated with two sudden noise stimuli per trial. For
each group, the second stimulus occurred at different time
intervals after the first stimulus, namely, at 1, 2, 3, and 4 seconds
respectively, occurring apparently, at least for groups A, B, and
C, within the relative refractory phase. For each subject 200
trials spaced over twenty consecutive days were run, 10 trials
being run each day at 30- second intervals. All recording was
kymographic, a silently operating tuning fork record giving the
time in units of 0.01 second. For the 1- second interval subjects,
the experiment was continued for 300 trials. From the data
obtained, the following points of significance were noted.
1. A pronounced tendency to complete negative adaptation
of the components of the body-reflex of the guinea pig to sudden
noise occurred after a sufficient number of stimulations.
2. Amount and rate of negative adaptation was greatest in
the first 40 trials.
3. For the two responses recorded in each trial there were
16 LOUIS H. COHEN

found relatively great differences between individuals in the

amount and rate of negative adaptation of each response.
4. Stimuli falling in an apparently supernormal phase were
least effective in producing negative adaptation as shown by one
case.
5. Individual differences in the length of the relative refractory
phase of a given reflex seem to exist.
6. The earlier the second stimulus occurs in relative refractory
phase, the less effective were both the first and the second stimuli
in producing negative adaptation. Conversely, the later the sec-
ond stimulus occurs in relative refractory phase, the more effec-
tive were both stimuli in producing negative adaptation.
Although these data are limited in number, these generalized
conclusions seem warranted by the consistency of the relationships
observed. Further investigation with larger groups at various
points of the refractory phase, absolute and relative, and also
within the rebound, seems desirable.
These experimental data confirm the tentative conclusions of
Dodge and Louttit and further permit the assumption of a
quantitative direct relationship between rate of adaptation of
reflex response and the interval between the first and the second
stimulus within relative refractory phase. Each future investiga-
tion concerned with the problem of reflex adaptation, therefore,
may well involve, whenever possible, a determination of the
refractory phase and rebound of the reaction system studied in
order that more complete knowledge of the relative excitability
of the system and the consequent susceptibility of the response to
modification may be obtained.