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Priming-produced facilitation or diminution of responding to

a Pavlovian unconditioned stimulus

Article  in  Journal of Experimental Psychology Animal Behavior Processes · November 1981

DOI: 10.1037//0097-7403.7.4.295 · Source: PubMed

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 Journal of Experimental Psychology:
Animal Behavior Processes
VOL. 7, No. 4 OCTOBER 1981

Priming-Produced Facilitation or Diminution of Responding

to a Pavlovian Unconditioned Stimulus
Nelson H. Donegan
Yale University
Two experiments, concerned with signal-produced variation of unconditioned
response (UR) amplitude, evaluated the roles of US intensity and response mea-
sure in determining when signaling the US results in a conditioned diminution
or a conditioned facilitation of responding. In Experiment 1, rabbits received
discrimination training, in an eye-blink-conditioning preparation, with a 1-mA,
50-msec shock US. In testing, preceding the US by CS+ facilitated the eye blink
response compared with preceding the US by CS~ or neither CS. In Experiment
2, subjects received discrimination training with a 5-mA US and were tested
with 1-, 2-, and 5-mA USs. During testing, subjects' eye blink responses (for
which robust CRs were observed) and gross body movement responses (for which
no CRs were observed) were simultaneously recorded. For the eye blink response,
preceding the US by CS+ facilitated responding to the 1-mA US, produced
negligible differences in responding to the 2-mA US, and diminished responding
to the 5-mA US, compared with preceding the US by CS~ or neither CS. For
the movement response, preceding the US by CS+ diminished responding at all
three US intensities, with the decremental effects increasing with increases in
US intensity. The results of Experiments 1 and 2 are discussed in terms of the
dual effects of CS+: the dimnution of US processing and the contribution of the
conditioned response to the measured response.

Considerable research in Pavlovian con- measures as a result of recent presentation

ditioning suggests that a target stimulus may of the same, or of an associatively related,
be rendered less effective in a variety of stimulus (e.g., Kamin, 1969; Terry, 1976;
Terry & Wagner, 1975; Wagner, Rudy, &
This article is based on a dissertation submitted to Whitlow, 1973). To approach such findings,
Yale University in partial fulfillment of the require- Wagner (e.g., 1976, 1978) proposed that a
ments for the PhD degree. The research reported herein .._f , .7, , , iji,_i,, <-„ ~A™*;nn o«
was supported in part by National Science Foundation Stimulus Will be less ikely to occasion an
GrantBNS77-16886toAllanR.Wagner.IthankAllan active representational state of rehearsal
R. Wagner for his continuous encouragement and many when it is prerepresented (primed) in short-
helpful suggestions throughout the course of research term memory (STM)
SbeJTSr I'A™S IIL^ The present studies are concerned with the
merous helpful comments in their role as members of applicability of this reasoning to Slgnal-pro-
my advisory committee, and Robert O. Crowder and duced variation in the immediate response
Michael Davis for their helpful comments in their role to an unconditioned Stimulus (US). Wagner
as readers on my committee (
v 1 9 7 6 ) suggested
ee that the response
r
decre-
Requests for reprints should be sent to Nelson H. ' . „. , ., .. „ , ,. .,
Donegan, who is now at the Department of Psychology, ment during habituation may be attnb-
Stanford University, Stanford, California 94305. uted to a transient priming in STM by recent

Copyright 1981 by the American Psychological Association, Inc. 0097-7403/81/0704-0295S00.75

295
296 NELSON H. DONEGAN
exposure to the same stimulus and/or a more
persistent priming in STM by contextual
cues to which the stimulus has become as-
sociated. In support of the likelihood of the
latter influence, Wagner (1976) pointed to
the so-called conditioned diminution of the
unconditioned response (UR; e.g., Kimble
& Ost, 1961; Kimmel, 1966) in which UR
amplitude has been reported to be reduced
as a specific result of the US being preceded
by an associated conditioned stimulus (CS).
A problem for this account is that a con-
ditioned diminution of the UR has not been
uniformly observed. Particularly trouble-
some are the reports of Grevert and Moore
(1970) and Hupka, Kwaterski, and Moore
(1970). Using a nictitating-membrane-con-
ditioning preparation in the rabbit, similar
to the preparation providing much of the
existing support for the decremental effect
of priming (e.g., Terry, 1976; Terry & Wag-
ner, 1975; Wagner, Rudy, & Whitlow,
1973), Grevert and Moore, and Hupka et
al. found that UR amplitudes were larger
on CS—>US trials than on US-alone trials.
These findings lead Hupka et al. to suggest
that whereas a conditioned diminution of the
UR may be characteristic of human eye
blink conditioning, it is not a similar feature
of conditioning in the rabbit.
Before accepting the proposal that a con-
ditioned diminution of the UR is not a phe-
nomenon of rabbit eye blink conditioning,
and its negative implications for Wagner's
priming theory, one should be cautioned by
two observations. First, it is known (e.g.,
Ison & Leonard, 1971; Young, Cegavske,
& Thompson, 1976) that a stimulus that
might be employed as a CS can have un-
learned reflex modulatory influences on a
shortly following US. The studies of Grevert
and Moore (1970) and Hupka et al. (1970)
involved few assurances that the greater re-
sponse on CS—>US than on US-alone oc-
casions was attributable to associative prop-
erties of the CS, free of such nonassociative
effects. Second, as Wagner (1979) noted, the
response to the US per se may be obscured
by the subject's conditioned response to the
CS that persists into the measurement pe-
riod; that is, a CS may have two associative
effects on responding at the time of the US,
one attributable to the CR that is generated,
the other attributable to a diminished pro-
cessing of the US. Whether one observes
evidence of a conditioned diminution of the
UR may depend upon the combination and
relative balance of these two effects.
Experiment 1 was primarily designed to
evaluate the reproducibility of the "UR" fa-
cilitation effect reported by Grevert and
Moore (1970) and Hupka et al. (1970) in
rabbit eye blink conditioning, under condi-
tions that more stringently assessed its as-
sociative basis. The facilitating effect was
observed. Experiment 2 was consequently
designed to comment on the argument that
facilitation or diminution may be variously
obtained, depending upon the combination
of CR and UR to the response measure. It
included the measurement of gross body
movement as well as the eye blink response
and provided, in comparisons with Experi-
ment 1 and in internal tests, an evaluation
of one parameter (the intensity of the test
US) that should influence the relative con-
tribution of the CR and UR. The findings
support the view that an associatively sig-
naled US produces a diminished UR.
Experiment 1
Given the reports of UR modulation by
associatively neutral stimuli (e.g., Hoffman
& Ison, 1980), there is reason to be cautious
in assuming that the CS-produced facilita-
tion reported by Grevert and Moore (1970)
and Hupka et al. (1970) had an associative
basis. Young et al. (1976) recently docu-
mented an unconditioned facilitation of the
nictitating membrane response by a tonal
stimulus that preceded an air puff US at a
temporal interval approximating that em-
ployed by Hupka et al. Consistent with this
finding is the fact that Grevert and Moore
observed greater responding on CS—.US
trials, relative to US-alone trials, at the out-
set of training. The only reassuring data pro-
vided by Grevert and Moore and by Hupka
et al. were that the facilitating effects in-
creased during the course of training. While
such observations are consistent with the
assumption that the CS became more effec-
 UR MODIFICATION BY CS PRIMING
tive as its association with the US increased,
they are not strong assurances.
In the present experiment, the associative
basis for a CS modification of the UR was
assessed in two ways. The first evaluated the
importance of the CS—>US correlation. Af-
ter initial discrimination training in which
one CS (CS+) was consistently followed by
a US and another CS (CS~) was not fol-
lowed by the US (except on infrequent probe
test trials), test sessions were administered
in which the US was preceded by CS+, CS",
or neither CS. Then subjects were given re-
versal training to determine whether the
ordering of response amplitude to the par-
ticular CS—»US combinations would be
reversed relative to the orderings observed
following the initial acquisition phase. The
second evaluation involved assessment of the
degree to which the effects of the CS might
be time locked to the CS—>US interval em-
ployed during training. A number of obser-
vations noted in the literature indicate that
a conditioned response to a CS occurs at the-
time when the US typically occurs (Hoehler
& Leonard, 1976; Millenson, Kehoe, & Gor-
mezano, 1977; Sears, Baker, & Frey, 1979).
If subjects' responding to a US is modulated
by learned response tendencies under control
of the CS and if the latter are timed to occur
at the time the US normally occurs, then the
effects of preceding a US by the CS might
vary as the time between CS and US onsets
deviates from the training interval. This pos-
sibility was evaluated in the last phase of the
experiment by running a series of test ses-
sions in which CS duration prior to US onset
was varied.
Method
Subjects. The subjects were eight male New Zea-
land white rabbits weighing between 2 and 3 kg at the
start of training. Each subject was housed individually
and maintained with ad lib food and water except during
experimental sessions.
Apparatus. During training and testing, each subject
was loosely held in a 51 X 18 X 14 cm Plexiglas box,
through which only its head protruded. The restraining
box was located in a 66 X 64 X 53 cm isolation chamber,
dimly illuminated by a 15-W neon bulb and maintained
with white noise at 64 dB (re 20 ,uN/m 2 ) measured at
the position of the subject's head (General Radio Sound
Pressure Meter No. 1551-C, A scale).
297
Two conditioned stimuli were used in the present
study. An auditory conditioned stimulus, a 3500-Hz
tone, was presented through the same speaker as the
white noise (the speaker was located behind the subject
on the rear wall of the chamber). The intensity of the
tone was approximately 10 dB over the background
noise level. A vibrotactile CS was presented by means
of a hand-massager (Valmour Model 880) mounted on
the floor of the restraining box so as to maintain firm
contact with the rabbit's chest. The total duration of the
CSs was 1,050 msec. The unconditioned stimulus was
a 50-msec train of 100/sec 1.0-mA square-wave shock
pulses, produced by an Argonaut LRA-046 constant-
current generator, and was delivered through stainless
steel electrodes (Sklar surgical wire, 32 ga.) sutured in
the skin about the orbit of the rabbit's right eye. One
electrode was implanted approximately 5 mm below the
extreme nasal extent of the eye; the other, approximately
5 mm above the extreme lateral extent. When presented
on reinforced trials, the US overlapped the last 50 msec
of the CS, which thereby determined a 1,000 msec
CS->US interval.
Closures of a subject's right eyelid were monitored
by a microtorque potentiometer which was taped to the
subject's head and communicated with the lid by a
length of thread hooked to a small permanent suture
loop. Movements of the eyelid turned a counterweighted
wheel affixed to the axle of the potentiometer. Resulting
resistance changes were graphically recorded on a Beck-
man Dynograph adjusted such that a 1-mm eyelid move-
ment produced a 2-mm deflection of the recording pen.
A conditioned response (CR) was defined as an eyelid
closure of .5 mm or more, requiring a graphic deflection
of at least 1 mm, during the interval 140-1,000
msec after CS onset. The amplitude of the eye blink
response at the time of the US occurrence was defined
as the maximum pen deflection, from the pre-CS base-
line, during a 175-msec period beginning with the US
onset.
Procedure. On the day prior to training, subjects
were introduced to the confinement conditions by being
placed in the restraining box outside the isolation cham-
ber for approximately 2 hr. During this period subjects'
heads were shaved and the shock electrodes were im-
planted.
The experimental design called for five distinct phases
of training. In the first phase, acquisition, all subjects
received eight daily sessions of Pavlovian discrimination
training with the auditory and vibratory CSs. Each ses-
sion consisted of 50 presentations of each CS, according
to an ABBA sequence, with intertrial intervals ranging
from 90 to 150 sec (M ITI = 120 sec). For half of the
subjects, the auditory CS was reinforced (CSA+), and
the vibratory CS was nonreinforced (CSB~). For the
remaining subjects, the vibratory CS was reinforced
(CSA+), and the auditory CS was nonreinforced (CSB~).
During each session five test trials were introduced on
which CSB was followed by the US. One test trial oc-
curred after each 20 discrimination training trials.
In the second phase of training (designated as Test
1), subjects were given two sessions of testing. These
sessions differed from the sessions in the first phase only
in that subjects also received five US-alone test trials
298 NELSON H. DONEGAN
during the course of discrimination training. In each
session one test trial (CSB followed by the US, or the
US alone) occurred after every 10 discrimination train-
ing trials. The ordering of test trials varied according
to a single alternation sequence, with the sequence being
reversed across sessions.
In the third phase, reversal training, subjects received
eight sessions of training in which the reinforcement
contingencies were reversed, i.e., CSB was reinforced
and CSA was nonreinforced. As in Phase 1, five test
trials, presentations of CSA followed by the US (written
herein as CSA~US), were introduced, one test trial oc-
curring after each 20 discrimination training trials.
In the fourth phase (designated as Test 2), subjects
received two sesssions of testing in which CSA~US and
US-alone test trials were presented such that one test
trial occurred after every 10 discrimination training
trials. The test trials varied according to a single alter-
nation sequence, with the sequence being reversed across
sessions.
In the fifth phase, the effect of varying the CS^US
interval on subjects' response to the US was assessed.
Subjects received 54 CSB+ and 54 CSA" discrimination
training trials, which occurred in an ABBA sequence,
in each of eight sessions. During each session three kinds
of test trials were introduced, one after every six training
trials. Each subject received six presentations of CSA
reinforced at a 1-sec CS—»US interval (time from CS
onset to US onset) and six presentations each of CSA
reinforced and CSB reinforced for which the CS—.US
interval was .5, 2, 4, or 8 sec in different sessions. The
CSA and CSB test trials in each session were presented
in a sequence that counterbalanced first-order sequential
dependencies. The successive test sessions included one
of the four intervals in first an ascending and then de-
scending order.
Results and Discussion
Discrimination and reversal training.
The effects of discrimination training on
subjects' conditioned responding to CSA and
CSB during acquisition and reversal training
can be seen in Figure 1 which presents the
mean CRs over all 100 CS+ trials and the
10 reinforced test trials with CS~ in each
two-session blocks of training trials. (The
CS~US test trials were used so as to include
only those trials for which UR measure-
ments were available. The data would be
essentially identical if based on the 50 CS~
training trials.) During the initial acquisition
phase, the mean percentage CRs to CSA
(CS+) increased across sessions, and the
mean percentage CRs to CSB (CS~) initially
increased and then decreased across sessions.
Only in the last block of sessions of this
phase was the mean percentage CRs to CSA
greater than the mean percentage CRs to
CSB. In reversal training, the mean per-
centage CRs to CSA (now, CS~) decreased,
and the mean percentage CRs to CSB (now,
CS+) increased across sessions. In the first
block of sessions, the mean percentage CRs
to CSA was greater than to CSB, as would
be expected on the basis of the reinforcement
contingencies during acquisition, whereas in
the last block of sessions the mean percent-
age CRs to CSB was greater than to CSA.
Results of statistical analyses of the initial
acquisition data indicated that there was a
reliable interaction of trial type (CSA vs.
CSB) with sessions, F(l, 42) = 5.26, p < .01.
Subsequent paired comparisons indicated
that the percentage CRs to CSA was not re-
liably different from that to CSB in the first
block of sessions, t(l) = 1.10, but was reli-
ably greater in the last block of sessions,
;(7) = 4.47, p < .01.
Results of statistical analyses of reversal
training indicated that there was again a
reliable interaction of trial type with ses-
sions, F(l, 42) = 12.78, p < .01. The per-
centage CRs to CSA was reliably greater
than the percentage CRs to CSB in the first
block of sessions, t(l) = 2.54, p < .05, but
was reliably less in the last block of sessions,
K7) = 4.34,p< .01.
Evaluation of responding in the US period
during initial acquisition training revealed
that the mean peak response amplitudes on
the 100 CSA+US and 10 CSB"US trials in
the first block of two sessions were 3.6 mm
and 3.8 mm, respectively, t(l) < 1.0, and in
the last block of two sessions were 6.3 mm
and 4.8 mm, respectively, t(l) = 3.59, p <
.01. The interaction of cuing condition (CSA
vs. CSB) with sessions was reliable, F(l,
42) = 4.58, p < .01. During reversal train-
ing, the mean peak response amplitudes in
the US period on the 10 CSA~US and
100 CSB+US trials in the first block of two
sessions were 6.0 mm and 5.4 mm, respec-
tively, t(l) = 2.25, p = .06, and in the last
block of two sessions were 5.0 mm and 6.7
mm, respectively, t(7) = 3.48, p < .05. The
interaction of cuing condition (CSA vs. CSB)
with sessions was reliable, F(l, 42) = 7.42,
/x.Ol.
Tests 1 and 2. The mean percentages
 UR MODIFICATION BY CS PRIMING
ACQUISITION
100
•-• CS A +
o-o CS B -
80
60
40
20
1 2 3 4
BLOCKS
Figure 1. Mean percentage eye blink CRs in acquisition, during which CSA served as CS+ and CSB
served as CS~, and in reversal training, during which CSB served as CS+ and CSA served as CS~, for
pairs of sessions in Experiment 1.
CRs to CSA and CSB across the two sessions
of Test 1 were 49.4 and 12.5, respectively,
F(l, 6) = 18.94, p < .01, and across the two
sessions of Test 2 were 13.8 and 66.5, re-
spectively, F(l, 6) = 55.94, p < .01.
Figure 2 shows the mean amplitude of
subjects' responses during the US period
over Test 1 and over Test 2 for the 100 trials
on which the US was preceded by CS+, the
10 trials on which the US was preceded by
CS~, and the 10 trials on which the US was
presented alone. The results of Test 1, in
which CSA served as CS+ and CSB as CS",
appear on the left-hand side of Figure 2. As
can be seen, preceding the US by CSA in-
creased the amplitude of responding relative
to US-alone presentations. Statistical anal-
yses revealed that the differences in response
amplitude among the three trial types (CSA,
CSB, and US alone) were reliable, F(2,
12) = 12.72, p < .01. The amplitude of re-
sponding on CSA trials was reliably greater
than on CSB or US-alone test trials,
t ( l ) = 3.40, p < .05 and t(l) = 4.12, p <
.01, respectively, and responding on CSB
trials was reliably greater than on US-alone
trials, t(l) = 2.75, p < .05.
The results of Test 2, in which CSB served
as CS+ and CSA served as CS~, appear on
299
REVERSAL
•-• CS A -
o-o CS B +
1 2 3 4
OF 2 SESSIONS
the right-hand side of Figure 2. As can be
seen, preceding the US by CSB facilitated
responding relative to preceding the US by
CSA, which in turn facilitated responding
relative to US-alone presentations. Statisti-
cal analyses revealed the differences in re-
sponse amplitude among the three trial types
(CSA, CSB, and US-alone) were reliable,
F(2,12) = 43.12, p < .01. Subsequent paired
comparisons indicated that the amplitude of
responding on CSB trials was reliably greater
than on CSA or US-alone test trials,
t(7) = 4.57,;? < .01 and t(l) = 8.69,p < .01,
respectively, and that responding on CSA
trials was reliably greater than on US-alone
trials, f(7) = 4.14, p < .01. In summary, the
same pattern of results was obtained in Tests
1 and 2; eye blink response amplitude at the
time of the US was greater when the US
was preceded by CS+ than when the US was
preceded by CS~ or neither CS.
The preceding results show that the CS+
(a) is capable of eliciting an observable eye
blink CR and (b) can increase the eye blink
response measured during the US period,
relative to CS~ or neither CS. In order to
assess the possible relation between these
two findings, the response amplitudes on the
CS~US and US-alone test trials, and on each
300 NELSON H. DONEGAN

TEST 1 TEST 2
12.5

E
J 10.0
ui
cc
13
U)
o 7.5
_J
o

5.0

2 2.5

CSB- NONE CSA- CSB+ NONE

CS PRECEDING THE US
Figure 2. Mean amplitude of the eye blink response at the time of US presentation when the US was
preceded by CS1, CS~, or neither CS in Test I (left-hand side) and Test 2 (right-hand side) of Experi-
ment I.

of the CS+US trials that immediately pre-
ceded a CS~US test trial, in the two sessions
of Test 2 were measured at periodic intervals
during the CS, US, and post-US periods.
Samples were taken at .25-sec intervals from
the time of CS onset, at .050-sec intervals
from the time of US onset to .25 sec after
US onset, and .5 and l.O sec after US onset.
Figure 3 shows the average response ampli-
tude at these various points in time on
CS+US, CS~US, and US-alone trials. As
would be expected on the basis of the level
of conditioned responding to CS+ and CS~,
there was a greater amount of eyelid closure
prior to US onset on CS+ than on CS~ or
US-alone trials. It is also apparent that the
peak response amplitude during the US pe-
riod was greatest on CS+US trials, at an in-

IO.O
•—• US PRECEDED B Y C S +
O—O US PRECEDED B Y C S -
7.5 D—Q US ALONE

5.0

-.75 -.50 -.25 0 .25 .50 l.O

SECONDS FROM US ONSET

Figure 3. Amplitude of the eye blink response prior to, during, and after presentation of the US when
the US was preceded by CS+, CS", or neither CS in Test 2 of Experiment I. (Each point represents
the mean amplitude of the eye blink response at the designated times from US onset. Points falling
between US onset and .25 sec after US onset are plotted at intervals of .05 sec.)
 UR MODIFICATION BY CS PRIMING
termediate level on CS US trials, and small-
est on US-alone trials. These data suggest
that the greater the level of responding prior
to US onset, the greater will be the likelihood
of observing facilitated responding in the US
period.
However, to say that the level of respond-
ing during the US period is related to the
level of responding during the CS period is
not to say that a detectable CR is a necessary
condition for observing facilitation. In Test
1, in which observable CRs occurred on ap-
proximately half of the CS+US trials, it was
possible to classify CS+ and CS" trials for
each subject as those on which a CR was
observed_(e.g., CR|CS + ) or not observed
(e.g., CRICS + ). A conditioned facilitation
of responding during the US period was
found in both types of CS+ trials. As ex-
pected, responding^ on CR | CS+ trials was
greater than on CR | CS+ trials, 7.4 mm and
6.1 mm, respectively. However, response
amplitude on CR | CS+ trials (6.1 mm) was
greater than on CR|CS~ trials (4.5 mm) in
seven of the eight subjects, and greater than
on US-alone trials (3.9 mm) in each of the
eight subjects. Hupka et al. (1970) also re-
ported facilitation of responding on CS+US,
relative to US-alone, trials prior to the de-
velopment of observable CRs. The present
findings of greater responding on CS+US
trials relative to CS~US trials in the absence
of observable CRs more clearly attest to the
associative basis of the facilitation. Wagner,
Thomas, and Norton (1967) interpreted
similar evidence of facilitation in the absence
of observable CRs as evidence for subthres-
hold tendencies of CS+ to provoke a response
that mimicked the UR, that could be de-
tected in the enhanced response to CS+US.
CS—>US-interval variation. Figure 4
shows subjects' mean eye blink response am-
plitude during the US period as computed
from all appropriate CS+US and CS~US
test trials presented in the CS—»US-interval
test sessions. (The line depicting response
amplitude on US-alone trials is the mean
response amplitude on US-alone trials oc-
curring in Tests 1 and 2. It is meant to pro-
vide an estimate of subjects' responding to
a US alone that can be compared with the
amplitude of the response to the US on
301
•—• US PRECEDED BY CS -
0—0 US PRECEDED BY CS -
10 0 US ALONE
in
O 75
O
O
UJ 5.0
UJ
5
5 I 0 2.0 4.0 80
CS-US ONSET INTERVAL (sec)
Figure 4. Mean amplitude of the eye blink response at
the time of US presentation when the US was preceded
by CS+ or CS~ at CS^US onset intervals ranging from
.5 to 8 sec in Experiment 1. (The dashed line represents
the mean amplitude of the eye blink response on US-
alone trials from Tests 1 and 2.)
CS+US and CS~US test trials at the various
CS—.US intervals.) Again, the amplitude of
responding at the time of the US was greater
when it was preceded by CS+ than by CS~.
Facilitation produced by CS+ was seen at all
of the CS—>US intervals and was greatest
at the 1-sec training interval.
Statistical analyses revealed that the dif-
ferences in response amplitude between
CS+US and CS~US trials was reliable, F(l,
6) = 45.50, p < .01, and that the interaction
of cuing conditon (CS+ vs. CS~) with the
CS—>US interval was also reliable, F(4,
24) = 5.05, p < .01. For further evaluating
the interaction, responding on CS+US and
CS~US trials at the 1-sec training interval
was compared with the responding on CS+US
and CS~US trials averaged across the re-
maining, nontraining, intervals (.5, 2, 4, and
8 sec). The mean response amplitudes during
the US period on CS+US and CS~US trials
having a 1-sec CS—>US interval were 7.0
mm and 4.7 mm, respectively, and for the
nontraining intervals were 5.4 mm and 4.3
mm, respectively. The interaction of cuing
condition with CS—»US interval was reli-
able, F(\, 6) = 36.61, p <. 01, and paired
comparisons indicated that responding on
CS+US trials was reliably greater than on
CS'US trials at the 1-sec training interval,
302 NELSON H. DONEGAN
•-• CS +
o-o CS-
i io.o
or
g 75
3
o trained at .5-, 1-, and 2-sec CS—»US inter-
D
^ 5.0
Ul
UJ
1 25
UJ
5 cally occurs, then each group should show
5 1.0 20 4.0 80
SECONDS FROM CS ONSET
5. Waveform of the eye blink CR to CS+ and
CS", estimated from trials on which the CR—>US onset
interval was 8 sec in tests of CS—>US onset variation
in Experiment 1. (CR amplitude was defined as the
maximum extent of eyelid closure during a 175-msec
window beginning .5, 1, 2, or 4 sec from CS onset. The
points at the 8-sec interval represent the mean amplitude
of the eye blink response 8 sec after CS onset.)
?(7) = 6.86, p<.0l, and across the non-
training intervals, t(l) = 6.18, p < .01.
One possible interpretation of the inter-
action of cuing condition with the CS—>US
interval is that the contribution of the CR
to the responding measured in the US period
is greatest at the time the US normally oc-
curs. By selecting CS—>US-interval test trials
having an 8-sec CS—>US interval, it was
possible to estimate the amplitude of the CR
at the time when the US was presented at
the training and several test intervals. To do
so, the maximum amplitude of subjects' re-
sponse was determined within a .175-sec
window beginning .5, 1, 2, and 4 sec from
the onset of CS+ and CS" as well as the
amplitude of the response at 8 sec after CS
onset. Figure 5 presents the average wave-
form of the CR on CS+ and CS~ trials. As
can be seen, the difference in response am-
plitude between CS+ and CS~ trials is great-
est at the 1-sec training interval relative to
the .5-, 2-, 4-, or 8-sec test intervals. These
observations are consistent with the proposal
that responding during the US period on
CS+US trials was greatest at the 1-sec in-
terval because the contribution of the CR
was greatest at that interval. An alternative
interpretation of the CS—>US-interval-func-
tion data presented in Figure 4 is that the
1-sec interval may be the optimal interval
for detecting the facilitating effects of pre-
ceding the US by CS+, independently of the
training interval. If groups of subjects were
vals and tested at each interval, one might
see the greatest facilitation at 1 sec for all
three groups. However, if facilitation is
greatest at the time at which the US typi-
maximum facilitation at the training inter-
val. If both factors influence response pro-
duction, one would expect to see greater fa-
cilitation at both the 1-sec and the training
intervals, relative to other nontraining inter-
vals.
In summary, the data reported in the pres-
ent experiment are consistent with the find-
ings of Hupka et al. (1970), that the am-
plitude of subjects' response on CS—.US
trials is greater than on US-alone trials. That
the greater response amplitude on CS+ trials
had an associative basis was demonstrated
by the findings that preceding the US by
CS+ resulted in larger responses than did
preceding the US by CS", after both original
training and reversal. Consistent with such
an associative interpretation are the findings
that the magnitude of the facilitation effect
was greatest at the CS—»US interval em-
ployed during training. These two sets of
data suggest that facilitation is greatest
when the US should be most expected on the
basis of the available cues, i.e., at a time
when the CR is likely to make its greatest
contribution to responding.
Experiment 2
In addressing reports of conditioned fa-
cilitation of the UR, Wagner (1979) pro-
posed that priming a target stimulus dimin-
ishes stimulus processing but that behavior
sequences initiated by the priming stimulus
may persist to summate with responding
generated by the target and thus make the
processing of the latter only appear to be
facilitated. With regard to the likely results
of CS—>US pairings when the CR that de-
velops to the CS mimicks the UR to the US,
 UR MODIFICATION BY CS PRIMING
Wagner (1979) noted that "even though US
processing (and thus UR magnitude) [are]
generally diminished when the stimulus [is]
associatively primed in STM by the CS, just
the opposite might appear to be the case
when the CS provokes a CR that mimicks
the UR and what is measured at the time
of the US is a combination of CR and UR"
(p. 77). This proposal, that, on a CS+US
trial, the measure of responding in a US
period is a combination of a conditioned re-
sponse and a diminished unconditioned re-
sponse (CS+US-^CR + UR'), suggests the
following generalization: The greater the
mimicking CR in relation to the UR, the
greater the likelihood of observing facilita-
tion on CS+US compared with US-alone
trials. Conversely, when a mimicking CR is
small in relation to the UR, one should be
more likely to observe a diminished UR on
CS+US relative to US-alone trials.
One experimental parameter likely to in-
fluence the ratio of the CR amplitude to UR
amplitude is the intensity of the US. The
lower the intensity of the US on occasions
of testing, the greater should be the ratio of
CR amplitude to UR amplitude, and the
greater should be the opportunity for ob-
serving facilitation. Support for such a pro-
posal can be found in the work of Hupka et
al. (1970) in which facilitation on CS+US
compared with US-alone trials was greater
when the US intensity was 2 mA than when
it was 4 mA. Also, Wagner et al. (1967),
investigating limb flexion conditioning in the
dog with a cortical US, found that preceding
a threshold value of the US by CS+ increased
the probability of a detectable UR but that
preceding a suprathreshold, training, value
of the US by CS+ diminished the amplitude
of responding, relative to US-alone presen-
tation.
The primary objective of Experiment 2
was to evaluate the effects of US intensity
on subjects' responding to signaled and un-
signaled USs. Subjects were trained and
tested as in Experiment 1, with the exception
that a higher, 5-mA, US was used rather
than the 1-mA US used in Experiment 1.
Consistent with the report of Wagner et al.
(1967), a conditioned diminution of the UR
was observed with the high-intensity US. A
303
series of test sessions was then run to eval-
uate whether conditioned facilitation and
conditioned diminution could each be ob-
served, depending upon the intensity of the
US during testing, independently of any dif-
ferences in the intensity of the US during
training that distinguished Experiments 1
and 2.
The secondary question in Experiment 2
concerned the possibility that the conse-
quences of preceding the US by CS+ depend,
in part, upon the response measure. It should
be recognized that the choice of response
measures provides another way of varying
the ratio of CR amplitude to UR amplitude
and, consequently, the likelihood of observ-
ing a conditioned diminution or a condi-
tioned facilitation of responding (e.g., by
selecting one response measure for which
robust CRs are observed and a second re-
sponse measure for which minimal or no
mimicking CRs are observed). In the present
experiment, the effects of a CS—>US se-
quence, versus US-alone presentation, were
evaluated with two response measures, the
eye blink measure, as employed in Experi-
ment 1, and a measure of gross body move-
ment. The eye blink measure, as has been
shown, exhibits an anticipatory CR that
mimicks the closure UR. The gross move-
ment measure, in contrast, revealed no ap-
parent anticipatory CR similar to the move-
ment produced by the US. As a result, it was
possible to evaluate the degree of condi-
tioned diminution and/or facilitation that
might be observed under conditions of test-
ing in which CS+ produced substantial mim-
icking CRs (eyeblink) or no apparent mim-
icking CRs (movement).
Method
The subject population, apparatus, and procedures
were identical to those of Experiment 1 except as noted.
Subjects. The subjects were eight naive male rabbits.
Apparatus. The apparatus was modified so as to
allow measurement of subjects' gross body movement.
An Astatic Model 16 phonograph cartridge was mounted
on the bottom of a 12.5 X 55.0 X 1.0 cm Plexiglas floor
that rested on two steel rods located at the front and
rear and 4.5 cm above the bottom of the restraining box.
The phonograph cartridge was located in the center and
304 NELSON H. DONEGAN
19 cm from the front end of the elevated floor. A metal
rod 1 mm in diameter and 50 mm in length was inserted
in place of a phonograph needle. Sudden movement on
the Plexiglas floor caused the metal rod to oscillate,
which in turn resulted in the phonograph cartridge pro-
ducing a voltage proportional to the displacement of the
metal rod. Weights were attached to the metal rods as
necessary to adjust the movement transducers such that
each was equally sensitive to a calibration procedure of
dropping a 50-g weight on the restraining box floor from
a distance of 6 cm.
The voltage output from the movement transducer
was graphically recorded by means of a Beckman Dy-
nograph with a gain setting of .5 V/cm. Any sudden
body movement of a subject resulted in the movement
transducer's producing an oscillating pattern of positive
and negative voltages which were graphically recorded
as an oscillating pattern of upward and downward pen
deflections from a pre-CS baseline level. The amplitude
of subjects' movement response was denned as
log(millimeter pen deflection + 1), where millimeter pen
deflection was determined by measuring the distance
between the maximum upward and downward pen de-
flections from the pre-CS baseline, in the .25-sec period
beginning with US onset.1 The amplitude of the eye
blink response at the time of US occurrence was defined
as the maximum pen deflection, from the pre-CS base-
line, during the same period. An interrupted light CS
produced by a Knight KG-323 strobe lamp located be-
hind the subject was substituted for the vibratory CS,
as the latter would itself have activated the movement
sensor. The US intensity on CS+ training trials was
5 mA.
Procedure. All rabbits received four sessions of dis-
crimination training, one session occurring per day. For
half of the subjects, the tone CS served as CS+ and the
light served as CS"; for the remaining half of the subjects,
the light served as CS+ and the tone as CS~. Within each
session subjects received 50 CS+ and 50 CS" trials. All
subjects received CS+ and CS" trials in the following
sequence, which was repeated five times during the course
of the session: +-+—++—++ +++—+. Dur-
ing discrimination training, only subjects' eye blink re-
sponses were recorded.
In each of the remaining sessions, subjects received
the basic discrimination training sequence described
above, during which additional test trials were period-
ically introduced. Following the initial acquisition phase,
subjects received a test session in which the discrimi-
native trials were supplemented with five presentations
of the US alone and five presentations of the US pre-
ceded by CS~, the test trials occurring in an alternating
sequence. The test trials were introduced after every
eight discrimination training trials, starting 20 trials into
the session. On each trial, subjects' movement and eye
blink responses were recorded simultaneously. After
testing, subjects received two sessions of discrimination
training.
In a subsequent series of test sessions, amplitudes of
responses were assessed to three different US intensities
when the US was preceded by CS+, CS", or neither CS.
In each test session, nine types of test trials were intro-
duced: trials on which a US of 1, 2, or 5 mA was pre-
ceded by CS+, CS", or neither CS. Each type of test
trial occurred three times during a test session, once in
each of three blocks of nine test trials. Each subject
received a different sequencing of the test trials, and in
each sequence the first-order transition probabilities
were approximately equated. Test trials were introduced
after every three discrimination training trials with the
usual, 5-mA US intensity, starting at the 20th training
trial of the session. Subjects received three such test
sessions, separated by two sessions of discrimination
training.
Following two sessions of further discrimination train-
ing, subjects were given two final test sessions in which
the CS—>US interval was varied in a manner similar to
that in Experiment 1. In each test session, subjects had
the 5-mA US preceded by CS+ or CS" at CS—US
intervals of .5, 1 , 2 , 4 , and 8 sec, or they had the US
presented alone. Hence, subjects received 11 types of
test trials, each type occurring three times per session
for a total of 33 test trials per session. A test trial oc-
curred after every three training trials, starting 20 trials
into the discrimination training sequence. Each type of
test trial occurred once in each of three blocks of 11 test
trials, and each subject received a different sequencing
of the test trials. Between the two test sessions, subjects
again received 2 days of discrimination training.
One subject in the group for which the tone served
as CS"1" and the light served as CS" died after the tests
in which the US intensity was varied. The effects of
varying the CS—.US intervals were determined with the
remaining seven subjects.
Results and Discussion
Discrimination training. The mean per-
centage eye blink CRs to CS+ and CS~ dur-
ing the course of discrimination training can
be seen in Figure 6. As is apparent, there
was a gradual development of greater re-
sponding to CS+. Statistical analyses of the
data from discrimination training indicated
that there was a reliable interaction of cuing
condition (CS+ vs. CS~) with sessions, F(3,
18) = 13.46, p < .01. The difference in per-
centage CRs between CS+ and CS" was not
reliable during the first session, F(l, 6) =
1.64, p > . 10, but was reliable during the last
session, F(l, 6) = 9.54, p < .05. There was
also a reliable interaction of cuing condition
(CS+ vs. CS~) with cue designation (tone
= CS+ vs. light = CS+), F(l, 6) = 28.32,
p < .01. As will otherwise be noted, the vi-
1
The movement response was defined as a log trans-
form of the millimeter-pen-deflection measure because,
with such a transform, the movement response grew in
a fashion more similar to the eye blink response as the
US intensity was increased.
 UR MODIFICATION BY CS PRIMING
sual CS was more likely to be followed by
an eye blink response than was the tonal
CS". Such responses to the visual CS" were
typically (but not always) of short latency
and were topographically discriminable from
CRs to the visual CS+. However, for lack
of consequences for the present purposes,
and for consistency with Experiment 1, no
attempt was made to further differentiate
the anticipatory closures.
Test 1. During Test 1, subjects continued
to respond differentially to CS+ and CS" in
terms of eye blink CRs (87.5% and 26.3%,
respectively), F(l, 6) = 15.16,p < .01. How-
ever, there was no detectable regular change
in the movement record as a consequence of
CS+ or CS" presentations, either in the form
of an episode of movement or in the form
of obvious quiescence. Movement was rarely
recorded except to the US, and when ap-
parent movement was observed during the
CS, it was almost always within a tracing
of less than 1 mm in amplitude that occurred
within .25 sec of CS onset. Because of the
lack of evidence of conditioned movement
changes, only the movement responses at the
time of US occurrence were subsequently
analyzed.
Figure 7 shows subjects' mean amplitude
eye blink response during the US period on
the 50 CS+US discrimination training trials
and the 5 CS~US and 5 US-alone test trials
100
80
60 -
z
UJ
o the US in each of the trial types of Test 1
£ 10
CL
<
UJ
2 3
SESSIONS
Figure 6. Mean percentage eye blink CRs to CS+ and
CS~ in each of the four initial sessions of discrimination
training in Experiment 2.
305
£
E 22.5
Ul
CC.
3
O 20.0
O
17.5
S 15.0
CS+ CS- NONE
CS PRECEDING THE US
Figure 7. Mean amplitude of the eye blink response at
the time of US presentation when the US was preceded
by CS+, CS~, or neither CS in Test 1 of Experiment 2.
in Test 1. As can be seen, the results are the
opposite of the results of Experiment 1; with
a 5-mA US, the response amplitude was
smallest on CS+US trials, at an intermediate
level on CS"US trials, and greatest on US-
alone trials.
Statistical analyses of the eye blink data
indicated that the differences among the
three trial types (CS+US, CS"US, and US-
alone) were reliable, F(2, 12) = 5.44, p <
.05. The response amplitude on CS+US trials
was reliably less than on US-alone trials,
t(l) = 3.01, p < .05, but the difference in
the response amplitudes between CS"US
and CS+US trials or CS"US and US-alone
trials did not reach statistical reliability,
t(l) = 1.69, p > .10 and t(l} = 1.40, p > .10,
respectively.
The topography of the eye blink response
prior to the US, during the US, and after
was determined as in Experiment 1. Figure
8 shows the average eye blink response am-
plitude at various points in time in relation
to US onset on CS+US, CS"US, and US-
alone trials. As would be expected on the
basis of the data presented in Figure 7, the
mean peak response amplitude following the
US on CS+US trials was smaller than on
CS~US or US-alone trials, even though the
mean amplitude of the eye blink response
306 NELSON H. DONEGAN

20.0
17 5 •—• US PRECEDED BY CS +
0-0 US PRECEDED BY C S -
15.0 a—o US ALONE

12.5
10.0

7.5
5.0

2.5
0
-I. -75 -.50 -.25 0 .25 50 l0
SECONDS FROM US ONSET

Figure 8. Amplitude of the eye blink response prior to, during, and after presentation of the US when
the US was preceded by CS+, CS~, or neither CS in Test 1 of Experiment 2. (Each point represents
the mean amplitude of the eye blink response at the designated times from US onset. Points falling
between US onset and .5 sec after US onset are plotted at intervals of .05 sec.)

immediately prior to the US onset was
greater on CS+US than on CS'US or US-
alone trials. The early increase and later
decrease in the mean eye blink response am-
plitude during the CS~ is almost entirely
accounted for by the tendency of the light,
when serving as CS~, to occasionally elicit
a short-latency response.
The effects of preceding the US by CS+,
CS~, or neither CS on the amplitude of sub-
jects' movement response at the time of US
occurrence are presented in Figure 9. Con-
sistent with the findings with the eye blink
response, the movement response amplitude
was smallest on CS+US trials and largest on
US-alone trials.
Statistical analyses of the movement re-
sponse data indicated that the differences in
movement response amplitudes among the
three trial types were reliable, F(2, 12) =
9.64, p<.0\. The response amplitude on

UJ
o:
UJ .2
o: t> o
(-5 3a _i
o

5 g •—• US PRECEDED B Y C S +
uj 5. 0—0 US PRECEDED B Y C S -
US ALONE

UJ
5
2
cs-us INTERVAL (sec)
CS + CS- NONE
CS PRECEDING THE US Figure 10. Mean amplitude of the eye blink response
at the time of US presentation when the US was pre-
Figure 9. Mean amplitude of the movement response ceded by CS+ or CS~ at CS^US onset intervals ranging
at the time of US presentation when the US was pre- from .5 to 8 sec in Experiment 2. (The dashed line rep-
ceded by CS+, CS~, or neither CS in Test 1 of Exper- resents the mean amplitude of the eye blink response
iment 2. on US-alone trials.)
 UR MODIFICATION BY CS PRIMING
CS+US trials was reliably less than on
CS~US and US-alone trials, *(7) = 3.01,
p < .05 and t(7) = 3.40,;? < .05, respectively.
The difference between CS~US and US-
alone trials was not reliable, t(l) - 1.48,
p> .10.
CS—.US-interval variation. Figure 10
shows subjects' mean eye blink response am-
plitude during the US period as computed
from all appropriate CS+US, CS~US, and
US-alone test trials presented in the
CS-^US-interval test sessions. The results
of varying the CS—^US interval are similar
to those of Experiment 1 in the sense that
the largest difference between CS+ and CS"
trials occurred at the 1 -sec training interval.
However, unlike in Experiment 1, and like
the results presented in Figure 7, the re-
sponse amplitude was smaller on CS+ trials
than on CS" or US-alone trials. The results
also differ from those of Experiment 1 in
that there was virtually no difference in re-
sponse amplitude between CS+ and CS~
trials at intervals other than the 1-sec train-
ing interval.
Statistical analyses of the eye blink re-
sponse amplitude data indicated that the in-
teraction of cuing condition (CS+ vs. CS~)
with CS—>US interval approached conven-
tional levels of significance, F(4, 20) = 2.72,
p < .06. In order to assess responding at the
training and nontraining CS-^US intervals,
response amplitudes on CS+US and CS~US
trials at the 1-sec training interval were con-
trasted with responding on CS+US and
CS~US trials averaged across the remaining
CS—.US intervals. At the 1 -sec interval, the
mean amplitudes of responding on CS+US
and CS~US trials were 20.6 mm and 22.5
mm, respectively, and across the remaining
CS->US intervals were 22.4 mm and 22.6
mm, respectively. The interaction of cuing
condition with CS—>US interval was reli-
able, F(l, 5) = 9.30, p < .05. At the 1-sec
training interval, responding on CS+US
trials was reliably less than on CS~US and
US-alone test trials, t(6) = 2.98, p < .05 and
?(6) = 2.78, p < .05, respectively, but the
difference in response amplitude between
CS~US and US-alone trials was not reliable,
t(6) < 1.0. There were no reliable differences
among the US-alone trials and the CS+US
307
uj.j
«!
z-6
3JS
UJ a
SI •—• US PRECEDED BY CS +
0—0 US PRECEDED 8Y CS -
US ALONE
.5 1.0 20 4.0 8.0
CS-US INTERVAL (sec)
Figure I I . Mean amplitude of the movement response
at the time of US presentation when the US was pre-
ceded by CS+ or CS~ at CS^US onset intervals ranging
from .5 to 8 sec in Experiment 2. (The dashed line rep-
resents the mean amplitude of the eye blink response
on US-alone trials.)
and CS~US trials over the remaining
CS-^US intervals.
The effect of varying the CS—>US interval
on the amplitude of the movement response
during the US period is shown in Figure 11.
As can be seen, the pattern of the movement
responding at different CS—>US intervals is
similar to the pattern of eye blink respond-
ing. Preceding the US with CS+ diminished
the amplitude of responding compared with
preceding the US with CS~ or neither CS,
and the greatest diminution occurred at the
1-sec training interval.
Statistical analyses of the movement re-
sponse amplitude data indicated that the in-
teraction of cuing condition (CS+ vs. CS")
with CS—»US interval was reliable, F(4,
20)=17.63,/><.01. The mean amplitudes
of the movement responses on CS+ and CS"
trials at the 1-sec interval were 1.21 and
1.69, respectively, and across the remaining
intervals were 1.65 and 1.70, respectively.
The interaction of cuing condition with
CS—US interval was reliable, F(l, 5) =
78.86, p < .01. At the 1-sec training interval,
the difference in responding on CS+US trials
was reliably less than on CS~US trials and
US-alone trials, t(6) = 6.42, p < .05 and
t(6) = 6.69, p < .01, respectively. The dif-
ference in response amplitude between CS~
and US-alone trials was not reliable, t(6) =
1.34, p> .10.
308 NELSON H. DONEGAN
12 5
5 1 0 2 0 4.0 8.0
SECONDS FROM CS ONSET
Figure 12. Waveform of the eye blink CR to CS+ and
CS~, estimated from trials on which the CS^US onset
interval was 8 sec in tests of CS—>US onset variation
in Experiment 2. (CR amplitude was defined as the
maximum extent of eyelid closure during a 250-msec
window beginning .5, 1, 2, or 4 sec from CS onset. The
points at the 8-sec interval represent the mean amplitude
of the eye blink response 8 sec after CS onset.)
In order to estimate the level of the eye
blink CR for the periods of time at which
the US was variously scheduled during the
CS—»US interval test sessions, the maximum
amplitude of subjects' response on the test
trials having an 8-sec CS—*US interval was
determined within a .25-sec window begin-
ning .5, 1, 2, and 4 sec from the onset of the
CS+ and CS" as well as the amplitude of the
response just prior to the US (8 sec after CS
onset). Figure 12 shows the mean response
amplitide at each of the above times on CS+
and CS~ trials. As can be seen, the difference
in response amplitude between CS+ and CS"
trials is greatest at the 1 -sec training interval
relative to the .5-, 2-, 4-, or 8-sec intervals.
The greater response amplitude on CS~ rel-
ative to CS+ trials at the .5-sec interval again
reflects the tendency of the light CS" to elicit
a short-latency eye blink response in the sub-
jects with that cue designation.
A particularly interesting feature of the
data presented in Figures 10 and 12 is that
on CS+US trials, at the 1-sec training in-
terval, the amplitude of subjects' response
during the US period was lower than in any
other test condition (see Figure 10), yet at
that corresponding moment the amplitude
of the CR was greater than in any other test
condition (see Figure 12).
Variation in US intensity. Across the
three test sessions in which the US intensity
was manipulated, subjects' mean percent-
ages eye blink CRs on CS+US and CSTUS
test trials were 89.4 and 44.4, respectively,
F(\, 6) = 38.91, p < .01.
The effects of preceding low-, intermedi-
ate-, and high-intensity USs by CS+, CS",
or neither CS on the amplitude of subjects'
eye blink responses during the US period,
averaged over the three test sessions, are
shown in Figure 13. As can be seen, with a
1-mA US the results are comparable with
those of Experiment 1: The amplitude of the
response on CS+ trials was greater than the
amplitude of the response on CS" and US-
alone trials. At the 5-mA intensity, the or-
dering of the trial types was reversed, rela-
tive to the ordering at the 1-mA intensity:
Response amplitude was smallest on CS+
trials, at an intermediate level on CS" trials,
and greatest on US-alone trials. Just as the
ordering -of trial types was reversed from
Experiment 1 (Figure 2) to Experiment 2
(Figure 7) when the training intensity of the
US was 1 and 5 mA, respectively, an iden-
tical reversal was seen within subjects when
•—• US PRECEDED BY CS +
°—° US PRECEDED BY CS -
n—n US ALONE
„ 20 0
E
CC 17.5
§
O
UJ
>-
UJ
UJ
5
1.0 20 6.0
US INTENSITY (mA)
Figure 13. Mean amplitude of the eye blink response
to 1-, 2-, and 5-mA USs when each US was preceded
by CS+, CS", or neither CS in Experiment 2.
 UR MODIFICATION BY CS PRIMING
the test intensity of the US was varied be-
tween 1 and 5 mA. At the intermediate US
intensity, 2 mA, the differences in the re-
sponse amplitude on different trial types
were less apparent.
Statistical analyses indicated that the dif-
ferences in eye blink response amplitude
among the three US intensities were reliable,
F(2, 12) = 69.48, p < .01; the higher the
shock intensity, the greater the response am-
plitude. The interaction of trial type (CS+US,
CS~US, and US-alone) with US intensity
(1, 2, and 5 mA), reflected by the reversal
in the orderings of the trial types at 1 and
5 mA, was reliable, F(4, 24) = 12.63, p <
.01. With the 1-mA US, the response am-
plitude on CS"1" trials was reliably greater
than on CS~ and US-alone trials, t(l) =
2.99, p < .05 and t(7) = 5.58, p < .01, re-
spectively, whereas the difference between
CS~ and US-alone trials was not reliable,
/(7) = 1.84,/» .10. For test trials on which
the 5-mA US occurred, the response ampli-
tude on CS+ trials was reliably less than on
CS~ and US-alone trials, t(l) = 2.68, p < .05
and ?(7) = 3.60, p < .01, respectively, and
the difference between CS~ and US-alone
trials was not reliable, t(l) = 1.75, p > .10.
The effects of preceding low-, intermedi-
ate-, and high-intensity USs by CS+, CS",
or neither CS on the amplitude of subjects'
•—• US PRECEDED BY CS ^
2.0 0—0 US PRECEDED BY CS -
UJ
CO
D—D US ALONE
CO
UJ
K.
US INTENSITY ( m A )
Figure 14. Mean amplitude of the movement response
to 1-, 2, and 5-mA USs when each US was preceded
by CS+, CS", or neither CS in Experiment 2.
309
movement responses during the US period
are shown in Figure 14. Preceding the US
by CS+ acted to decrease the amplitude of
responding at all US intensities, with dimi-
nution being greatest at the 5-mA US in-
tensity.
Statistical analyses indicated that the dif-
ferences in the movement response ampli-
tude among the test trials with the three US
intensities were reliable, F(2, 12)= 52.44,
p < .01, and that the differences in response
amplitude across test trial types (CS+US,
CS'US, and US-alone) were reliable, F(2,
12) = 7.32, p < .01. The interaction of trial
type with US intensity (1,2, and 5 mA) was
also reliable, F(4, 24) = 5.91, p < .01. With
the 5-mA US, the response amplitude on
CS+ trials was reliably less than on CS" and
US-alone trials, t(l) = 6.11, p < .01 and
t(7) = 5.78, p<.01, respectively, and re-
sponse amplitude on CS~ trials was reliably
lower than on US-alone trials, t(l) = 3.23,
p < .05. The differences among the various
trial types when the target US was 2 or 1
mA were not reliable.
The pattern of eye blink response data
resulting from variation in the US intensity
(Figure 13) indicates that preceding a low-
intensity US by an associatively related CS
produces increments in response amplitude
but that the same operations result in dec-
rements in response amplitude when the US
intensity is high. These findings support the
hypothesis made in the introduction that the
different effects of CS priming reported by
Kimble and Ost (1961), Hupka et al. (1970),
and Wagner et al. (1967) may reflect the
fact that the intensity of the target US de-
termines the occasions on which CS priming
will result in increments or decrements in
response amplitude. However, the compa-
rable pattern of movement response data
(Figure 14) indicates that US intensity is not
the sole determining variable, as preceding
the low-intensity US by CS+ did not result
in greater movement response amplitude
than did presentation of the US alone.
General Discussion
Contrary to the proposal of Hupka et al.
(1970), that a conditioned diminution of the
310 NELSON H. DONEGAN
UR is not a feature of conditioning in the
rabbit, the results of the preceding experi-
ments indicate that signaling the occurrence
of a US can result in a diminished UR (e.g.,
Figures 7, 9, 13, and 14) and are in agree-
ment with reports of a conditioned diminu-
tion of the UR in other species and response
measures (e.g., Kimble & Ost, 1961, Kim-
mel, 1966). The signal-produced decrements
in the eye blink and movement URs see in
Experiment 2 are clearly consistent with the
application of Wagner's priming theory to
the phenomena of "habituation," i.e., its as-
sumption that a primed US will be less well
processed and will generate a less vigorous
UR relative to presentation of the US when
not primed (Wagner, 1979). The rinding of
a conditioned diminution of the UR in Ex-
periment 2 is also in accord with numerous
experiments utilizing the rabbit eye blink
preparation that demonstrate priming-pro-
duced decrements in US processing in a va-
riety of alternative measures (e.g., Terry,
1976; Terry & Wagner, 1975; Wagner et
al., 1973).
The findings of a facilitated eye blink, but
not movement, response on CS+US trials
with the low-intensity US are consistent with
Wagner's (1979) proposal that responding
on CS+US trials is likely to be a combination
of the CR and a diminished UR. According
to this reasoning, the facilitation of the eye
blink response on CS+US trials was due to
the mimicking response tendency to the CS+,
whereas the absence of facilitation in the
movement response was due to the absence
of detectable movement CRs, i.e., CRs that
could be detected in the presence of CS+ or
in summation with the US. The fact that
facilitation was seen at the low, but not the
high, US intensity can be taken to indicate
that the relative contribution of a mimicking
CR to the response measure is greater when
the ratio of CR amplitude to UR amplitude
is high, as would be the case in the eye blink
when the test US is weak and elicits a
small UR.
It is interesting that a similar relation has
been observed in a variety of simple human
information-processing tasks, e.g., lexical
decision and naming tasks. In such tasks,
subjects' response to a target stimulus can
be considered UR-like in the sense that both
responses are relatively automatic and are
based on prompt recognition of the stimulus.
Subjects are typically required to make an
immediate response (e.g., naming) to a tar-
get stimulus that has been shortly preceded
by associatively related or unrelated stimuli.
The measure of principal interest is subjects'
reaction time, and the typical finding is that
reaction time is facilitated (reduced) when
the target is primed compared with when it
is not primed. Preceding a target word (e.g.,
butter) by an associatively related word
(e.g., bread), for example, results in shorter
reaction times in lexical decision and naming
tasks than if the target follows a nonrelated
word (e.g., iron; Becker & Killion, 1977;
Meyer, Schvaneveldt, & Ruddy, 1975;
Neely, 1977; Schvaneveldt & Meyer, 1973).
Consistent with the findings of Experiments
1 and 2, Hupka et al. (1970), and Wagner
et al. (1967), Becker and Killion (1977)
found that priming-produced facilitation in
lexical decision and naming tasks was great-
est at the lowest target-stimulus intensity
employed. Similar relations have been ob-
served in lexical decision tasks in which the
salience of the target stimulus was reduced
by embedding the letter string in a random-
dot array (Meyer et al., 1975) or rotating
the target word 180° (Massaro, Jones, Lip-
scomb, & Scholz, 1978). These correspon-
dences between the intensity of the target
stimulus and the likelihood that preceding
the target by an associated stimulus will re-
sult in a facilitated response suggest the pos-
sibility that models of performance devel-
oped to account for the priming effects seen
in the Pavlovian conditioning literature may
be applicable to the priming effects seen in
the literature on human information pro-
cessing and perception.
In addressing the issue of response gen-
eration, Wagner's priming theory (1976,
1979) assumes that the responding to a US,
and/or associated CS, seen in a variety of
response measures is mediated by the activ-
ity of a unitary US representation in STM.
That is to say, CS+ presentation produces
activation of the US representation and re-
sults, for example, in eye blink and move-
ment CRs that can be observed. Subsequent
 UR MODIFICATION BY CS PRIMING
presentation of the US results in diminished
activation of the US representation and, cor-
respondingly, diminished eye blink and
movement URs, relative to US-alone trials.
Differences observed in different response
measures, e.g., detectable eye blink CRs but
no detectable movement CRs, must then be
approached by assuming differences in the
mapping of US representational activity into
the different behaviors,
To suppose that different measures of re-
sponding mirror the same US processing in
STM and to allow for differences observed
across response measures in terms of differ-
ent response mapping functions is one thing,
but specifying the differences in the response
mapping is another. The general theoretical
challenge is to develop a model that clearly
specifies the consequences of CS+US and
US-alone trials on US processing and the
relations between changes in US processing
and changes in the response measures under
consideration. A formal characterization of
response generation that attempts to meet
these demands has been developed by Do-
negan and Wagner (Note 1) and has been
incorporated in a model of automatic mem-
ory processing (dubbed "SOP") recently
described by Wagner (1981). From the SOP
model's basic assumptions regarding stim-
ulus processing and response generation,
Donegan and Wagner developed theoretical
equations describing US representational
activity and the mapping of such activity into
responding on CS+US and US-alone trials,
appropriate to the test data depicted in Fig-
ures 13 and 14 from Experiment 2. It was
found that several equations well described
the qualitative and quantitative relations
observed within and across the eye blink and
movement response measures where the
equations for the two measures differed only
by a single parameter tied to the conse-
quences of associative activation, and a mul-
tiplicative constant.
Reference Note
I . Donegan, N. H., & Wagner, A. R. Conditioned dim-
inution and facilitation of the UR: A sometimes op-
ponent-process interpretation. Manuscript prepared
for publication in I. Gormezano, W. F. Prokasy, &
R. F. Thompson (Eds.). Classical conditioning III:
311
Behavioral, neurophysiological, and neurochemical
studies in the rabbit, book in preparation, 1981.
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Received December 29, 1980
Revision received June 3, 1981

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