Journal of Comparative and Physiological Psychology

1970, Vol. 72, No. 3, 492-497

EFFECTS OF STIMULUS INTENSITY AND INTERTRIAL

INTERVAL ON HABITUATION OF THE1 HEAD-SHAKE
RESPONSE IN THE RAT
HENRY R. ASKEW2
Michigan State University

The effects of stimulus intensity and intertrial interval on habituation and

retention of habituation of the head-shake response in the laboratory rat
was investigated using a 3 X 5 within-subjects factorial design. Both inde-
pendent variables had major effects on habituation of the head-shake re-
sponse. Theoretically, the most significant conclusion was that habituation
of the head-shake response appeared to be due to both a short-term dec-
remental process that occurred within single trials and took approximately
100 sec. to recover, and a long-term decremental process that occurred across
trials and showed only 70-80% recovery after a 30-min. rest period.

The present investigation was concerned
with the effects of stimulus intensity and
intertrial interval on habituation of the
head-shake response (HSR) to an air
stimulus in the laboratory rat. Previous
work by Askew, Leibrecht, and Ratner
(1969) and Leibrecht and Askew (1969)
have established the advantages of this
experimental situation for the study of
habituation.
Virtually all authors who have dealt with
habituation have observed that intensity
and ITI are major variables that affect the
phenomenon (e.g., Harris, 1943; Ratner &
Denny, in press; Thompson & Spencer,
1966). However, the specific effects of
these variables on habituation in different
stimulus-response systems have not always
been the same. With respect to stimulus
intensity, although higher intensities are
typically associated with higher initial
response levels and higher asymptotic re-
sponse levels (e.g., Davis & Wagner, 1968;
Dunlop, Webster, & Rodger, 1966; Kuen-
zer, 1958; Miller & Murray, 1966; Uno &
Grings, 1965), the results are less con-
sistent for other dependent variables. For
example, the amount of habituation is
sometimes reported to be greater (Dunlop
et al., 1966; Kuenzer, 1958; Miller &
1
This paper is based on a dissertation submitted
in partial fulfillment of the requirements for the
PhD degree, Michigan State University, 1969.
2
Requests for reprints should be sent to Henry
R. Askew, who is now at Fordham University,
Bronx, New York 10458.
492
Murray, 1966), sometimes less (Thompson
& Spencer, 1966; Uno & Grings, 1965),
and sometimes not different (Davis &
Wagner, 1968; Wolfensberger & O'Connor,
1965) the greater the stimulus intensity.
Similarly for the rate of habituation (num-
ber of trials necessary to reach asymptote),
although most workers found no difference,
two studies reported that weaker stimuli
habituate at a faster rate (Kuenzer, 1958;
Prechtl, 1958) and one study found that
the weaker stimulus took longer to habitu-
ate (Thompson & Spencer, 1966).
A similar lack of consistency has re-
sulted from studies dealing with the vari-
able of intertrial interval. Work with a
number of species has yielded evidence
that shorter ITIs resulted in a decrease to
a lower asymptotic response level than
longer ITIs with no difference in the rate
of decrement (Barrass, 1961; Brown,
1965; Schaub, 1965; Simons, Dunlop,
Webster, & Aikin, 1966; Winokur, Stewart,
Stern, & Pfeiffer, 1962). However, in other
cases (Prechtl, 1958; Prosser & Hunter,
1936) the difference produced by the dif-
ferent ITIs was in terms of the number of
trials necessary for the response level to
reach a zero level. In other words, there
was a difference in the rate of habituation
and no difference in the asymptotic level
of responding reached.
After reviewing the entire literature, it
is apparent that the variables of stimulus
intensity and ITI do not always have
 HABITUATION OF THE HEAD-SHAKE RESPONSE 493

predictable effects on habituation in dif- ter moved his hand accordingly to keep the stimu-
ferent stimulus-response systems. There- lus directly in the rat's ear. If the animal turned
around on the stand, the experimenter rotated the
fore, it appears to be necessary to look at stand until the rat was again in the proper orienta-
the effects of these two variables when a tion, at which time the normal stimulus pattern
new experimental situation is being stud- was resumed. The experimenter practiced during
ied. As a result, the present study was previous experiments (Askew et al., 1969; Lei-
designed to be a relatively thorough analy- brecht & Askew, 1969) and became relatively
proficient in maintaining the correct ear to test
sis of the effects of stimulus intensity and stimulus relationship for all but the most drastic
ITI on habituation and retention (or con- movements on the part of the rat. Distributed
versely, recovery) of habituation of the practice for an hour or two is probably sufficient
head-shake response (HSR). The fact to The reliably deliver the stimulus.
variables of ITI and stimulus intensity
that there are no appreciable carry-over ef- were manipulated in a 3 X 5 within-subjects fac-
fects after 24 hr. (Askew et al., 1969), torial design. The fixed ITIs of 1, 10, and 100 sec.
coupled with large but reliable individual were combined with stimulus intensities corre-
differences in the HSR level, led to the sponding to deflections of .5, 1, 2, 10, and 100 mm.
use of a within-subjects design. of the column of alcohol such that for each subject
on each session all trials with the air stimulus
were given with the same fixed ITI and intensity.
METHOD Therefore, each subject was given one session
Subjects corresponding to each of the 15 different combina-
tions of the two independent variables. The in-
The subjects were 15 male Holtzman albino rats terval between each of the 15 sessions was 24 hr.
85 days of age. All animals had been reared in with each subject being given the 15 experimental
group cages, had been fed and watered ad lib, and sessions in a randomly determined order.
were experimentally naive. Each session began with the rat being placed
on the stand and being allowed 5 min. to adapt to
Apparatus the stand. The animal was then given 10 15-sec.
Movement of the animals was restricted by trials (Trials 1-10) and then was immediately re-
placing them upon an elevated 2 X 6.5 in. platform turned to its home cage about 5 ft. away from the
(constructed of !/4-in. hardware cloth) 29 in. above test stand. After spending 25 min. in its home cage
the floor. Hanging downward and sloping outward the subject was returned to the stand for the re-
at an angle of 10° from the edges of the platform test phase, which consisted of another 5-min.
were sections of galvanized sheet metal, forming a adaptation period without air stimulation followed
collar 10 in. long to discourage escape attempts. by five additional 15-sec. trials (Trials 11-15) with
The base of the wooden column which supported the same ITI and intensity as was used for the
the platform was mounted on ball bearings so test phase of that particular session. This test-re-
that the entire test stand could be rotated in test design was used to investigate retention of
either direction. habituation.
A test stimulus consisted of pressurized air Twenty-four hours after the completion of the
from a Silent Giant aquarium pump (Model 120) last of the 15 experimental sessions, each subject
delivered through a hand-held rubber tube with an was given a base-rate control session consisting of
inside diameter of 1 mm. Adjustable valves were 10 15-sec. trials with a 10-sec. ITI and a "zero"
placed in the tube such that the air pressure at stimulus intensity. The tube was moved back and
the end of the tube could be precisely regulated. forth next to the ear, as in experimental sessions,
Pressure was measured by placing the end of the but the air stream was directed away from the
tube 1 mm. from the end of an open manometer head. This procedure resulted in auditory and
with an inside diameter of 1.5 mm. The manometer visual components of the test stimulus without the
contained enough alcohol to form 120-mm. columns air-pressure component. The HSRs were recorded
on both sides. This air system resulted in a range on this control session as on any other session.
of possible intensities corresponding to from a zero
to a 140-mm. deflection in the column of alcohol. Treatment of the Data
Procedure In order to handle possible complexities, the
habituation curves were analyzed in terms of
The HSR was elicited by moving the air stimu- several dependent variables which require defini-
lus back and forth across the center of the left ear tion. First, the initial HSR level referred to the
of the rat at an approximate rate of 3 ops. The dis- number of head-shake responses on the first trial.
tance of the tube from the ear was s/a—Vz in. and Similarly, the terminal HSR level referred to the
the locus of stimulation was approximately Ys in. number of head-shake responses on Trial 10. As
wide. Whenever the rat moved its head an appre- a measure of the relative amount of decrement the
ciable distance during stimulation, the experimen- percentage decrement was defined as the difference
 494 HENRY R. ASKEW

between the initial and terminal HSR levels di- eating that some retention of habituation
vided by the initial HSR level. The rate oj habitu-
ation was defined as the number of trials necessary did take place after the 30-min. retention
before the HSR level fell to its asymptotic re- interval.
sponse level (asymptotic response level was opera-
tionally defined as equal to the terminal level). Effects of Stimulus Intensity
Two measures of recovery were used. First, the Figure 1 gives the mean number of
amount of recovery was defined as the absolute HSRs per trial as a function of trial num-
difference in HSR level between Trials 10 and 11
(before and after the 30-min. retention interval). ber and stimulus intensity for the 10 test
Finally, as a measure of the relative amount of trials and the 5 retest trials. Looking first
recovery, the percentage recovery was defined as at the 10 test trials alone, there were highly
the amount of recovery divided by the amount of significant differences in the initial HSR
habituation (the absolute difference in HSR levels
between Trials 1 and 10 is defined as the amount level on Trial 1 (F = 71.5, df = 4/56,
of habituation). p < .01) and the terminal HSR level on
The data analyses consisted of scoring each sub- Trial 10 (F = 44.2, df = 4/56, p < .01).
ject's performance on each session in terms of the As can be seen from the figure, higher
several dependent variables discussed in the pre- intensities resulted in higher levels of re-
ceding section. Then each of the dependent varia-
bles was statistically analyzed using a three-way sponding both at the beginning and at
single observation per cell repeated-measures the end of the habituation session. With
analysis of variance (Winer, 1962, p. 290) with respect to the relative amount of habitua-
fixed factors of ITI and intensity and a random tion, although there was an overall sig-
factor corresponding to subjects.
nificant difference in the percentage decre-
RESULTS ment (F = 10.0, df = 4/56, p < .01),
this difference was due to the small per-
Before dealing with the effects of the centage decrease that occurred in the .5-
two independent variables, it is necessary mm. 'Condition which showed essentially no
to note that when all conditions were decrement. Without the .5-mm. condition
combined, the HSR level was significantly there was no significant difference in the
lower on Trial 10 than it was on Trial 1 percentage decrement among the other four
(t = 7.3, df = 14, p < .01), indicating intensities. Turning to the rate of habitua-
that habituation did, in fact, occur. Over tion, there was a highly significant dif-
all conditions, habituation involved a 41% ference among the intensities (F= 16.0,
decrement from the initial HSR level. df - 4/56, p < .01). Examination of Fig-
Turning to retention, the HSR level on ure 1 showed that higher intensities ap-
Trial 11 was significantly lower than on peared to take more trials for the HSR
Trial 1 (t = 6.0, df = 14, p < .01), indi- level to approach asymptote than did
lower intensities.
With respect to the effects of stimulus in-
tensity on the percentage recovery, there
was no significant difference among intensi-
ties.
Effects of Intertrial Interval
Figure 2 gives the mean number of
HSRs per trial as a function of trial num-
ber and intertrial interval. Although there
was no significant difference in the initial
HSR level, the difference in the terminal
15
HSR level was highly significant (F =
31.4, df = 2/28, p < .01). This result
FIG. 1. Mean number of HSRs per trial as a
indicated that the difference in ITIs was
function of trial number and stimulus intensity in terms of the asymptotic response level
(expressed as the number of millimeters of alcohol that occurred. There was no significant
displaced in a manometer). difference in the rate of habituation, with
 HABITUATION OF THE HEAD-SHAKE RESPONSE
z ITI, the within-trial habituation curves
2,
5 10
TRIAL NUMBER
FIG. 2. Mean number of HSRs per trial as a
function of trial number and intertrial interval.
the three conditions falling to the dif-
ferent asymptotes after approximately the
same number of trials.
Turning to recovery, there was a highly
significant difference among the three ITIs
in the amount of recovery. As can be seen
from Figure 2, there was only a very
slight increase from Trial 10 to Trial 11
for the 100-sec. ITI, but for the shorter
ITIs the amount of recovery was much
greater. It is also important to note that
there was no significant difference in the
number of HSRs on Trial 11 for the dif-
ferent ITIs. In conclusion, the major dif-
ference between the three ITI conditions
was that they produced different asymp-
totic response levels. Over the 30-min. re-
tention interval, however, these differ-
ences disappeared so that there was no
difference between the conditions on the
TRIAL 1 TRIAL 2 TRIAL
• I
1.0 O 10
• 100
w
tt
V)
.6
Z
<
.2
controk
1 3 5 1 3 5 1 3 5 1 3 5 1 3 5^r 1 3 5
3-SEC. WITHIN-TRIAL TIME PERIODS
FIG. 3. Mean number of HSRs per 3-sec. time period as a function of intertrial interval for several
selected trials.
495
first trial following the interval (Trial
11). This failure to find a difference after
the retention interval was not due to the
fact that all conditions recovered com-
pletely, since the overall percentage re-
covery was only about 75%.
In order to further analyze the effects of
for each ITI on several trials are presented
in Figure 3. Briefly, each 15-sec. trial was
divided into five 3-sec. periods and the
mean number of HSRs per 3-sec. time
15
period was then looked at for different
ITIs within each of the trials. From inspec-
tion of Figure 3 it seems clear that the
difference in HSR levels that occurred in
Figure 2 was due to the differential amounts
of recovery time between each trial cor-
responding to the different ITIs. Spe-
cifically, it appeared from Figure 3 that
with the 100-sec. ITI there was consider-
able trial-to-trial recovery with the HSR
level at the beginning of each trial being
relatively high. For the 1- and 10-sec.
ITIs, on the other hand, large amounts of
recovery between trials did not appear to
take place. It appeared, therefore, that
the significant difference between ITIs in
the asymptotic (or terminal) HSR level
shown in Figure 2 was a result of dif-
ferential recovery from trial to trial, with
different ITIs showing different response
levels during the first few seconds of each
trial (after Trial 1) but falling to the
same level by the last few seconds of each
trial. Apparently, a good deal of the
decrement which occurred within single
3 TRIAL 4 TRIAL 10
SEC.
SEC.
SEC.
496 HENRY R. ASKEW

trials was relatively short term, recovering Winokur et al. (1962), the difference in
quickly (within 100 sec.) after termina- asymptotes between the different ITIs ap-
tion of the stimulus. peared to be the result of allowing a
greater amount of recovery from short-
Interactions between Intensity and ITI term within-trial decremental effects
The only significant Intensity X ITI in- (short term in the sense that recovery
teraction was for the amount of recovery occurs by 100 sec.) the longer the time
(F = 4.2, df - 4/56, p < .01). Explana- between trials. In addition, a superim-
tion of this result lies in the finding that posed long-term decremental process also
for the 100-sec. condition, where there was has to be postulated. Evidence for the ex-
almost no recovery over the 30-min. in- istence of such a long-term effect is shown
terval, there was no difference among the by the substantial retention of habituation
intensities. across the 30-min. intersession interval.
Further support comes from a previous
DISCUSSION study of habituation of the HSR by
It appears that intensity did not greatly Leibrecht and Askew (1969) where habitu-
affect habituation per se, but rather in- ation was significantly retained for an in-
fluenced the habituation curves somewhat terval of at least 6 hr. In the present study,
indirectly by greatly affecting the overall the finding that all three ITIs recovered
level of responding. This conclusion does to approximately the same level after the
not agree with Thompson and Spencer's 30-min. intersession interval indicates that
(1966) characteristic of habituation deal- the variable of ITI did not appreciably
ing with intensity, which argues that affect the course of this long-term decre-
stronger stimuli show less habituation. mental process.
However, other studies with results some- A multiprocess interpretation of habitua-
what similar to the present study (Dunlop tion phenomena such as is employed in the
et al., 1966; Kuenzer, 1958; Miller & present study is not new. Hinde (1960)
Murray, 1966) seem also to argue against presents strong evidence that habituation
the universal applicability of Thompson of the mobbing response to an owl model in
and Spencer's intensity hypothesis. chaffinches is the result of the interaction
With respect to intertrial interval, the of both long-term and short-term incre-
findings that shorter ITIs were associated mental and decremental effects. A multi-
with a greater amount of habituation to a process interpretation also appears to be
lower asymptote with no accompanying supported by a consideration of the tre-
differences in the rate of habituation are mendous variability among the different
in line with the major portion of the litera- experimental situations in which habitua-
ture (e.g., Thompson & Spencer, 1966). tion has been studied, particularly in
The two major studies which do not fall terms of both (a) the degree of perma-
into this pattern (Prechtl, 1958; Prosser & nence of the decremental phenomena and
Hunter, 1936) appear to differ in that the (6) the underlying physiological mecha-
level of responding for all ITI conditions nisms which appear to be involved. It does
went to a zero level. In these cases it is not, for example, make much sense to argue
impossible to assess whether ITI had a that habituation of the orienting reflex,
completely different effect or whether some which has somewhat lasting effects and
sort of "floor" effect was operating to appears to be best explained on the basis
obscure the same relationship between ITI of central inhibition of afferent input, is
and habituation which is shown by the the result of the same underlying process
rest of the literature. as that occurring in Thompson and Spen-
The most theoretically important rinding cer's (1966) spinal cat preparation, where
of the present study is that at least two centrally mediated inhibition cannot be
processes appear to be necessary to ac- involved and complete recovery occurs
count for the effects of ITI on habitua- within 1 or 2 hr. For these reasons, it ap-
tion of the HSR. First, as suggested by pears most reasonable to take a somewhat
 HABITUATION OF THE HEAD-SHAKE RESPONSE 497

open-ended theoretical approach by ad- KUENZEK, P. P. Verhaltensphysiologische Unter-

mitting that there is really no reason to suchungen iiber das Zucken des Regerwurms.
Zeitschrift fur Tierpsychologie, 1958, 15, 31-49.
believe that the repeated application of a LEIBEECHT, B. C., & ASKEW, H. R. Habituation of
given stimulus cannot simultaneously ac- the head-shake response in the rat: Recovery,
tivate several underlying decremental transfer, and changes in topography. Journal
processes ranging from sensory adaptation, of Comparative and Physiological Psychology,
to some sort of altered synaptic transmis- MILLER,1969, 69, 699-708.
J. D., & MURRAY, F. S. Guinea pig's im-
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