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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

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 simultaneouslyrecorded. 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.

facilitated the eye blink

+

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, &

Whitlow, 1973). To approach such findings,

This article is based on a dissertation submitted to

Yale University in partial fulfillment of the require- Wagner (e.g., 1976, 1978) proposed that a

ikely to occasion an

GrantBNS77-16886toAllanR.Wagner.IthankAllan active representational state of rehearsal R. Wagner for hiscontinuous encouragement and many when it is prerepresented (primed) in short- helpful suggestions throughout the course of research term memory (STM)

ment s for th e Ph D degree. The research reported herei n was supported in part by National Science Foundation

f

,

.7, ,

Will be

iji,_i, , <-„ ~A™*; nn

Stimulus

,

less

SbeJTS r I'A™ S IIL ^

The present studies are concerned with the

merous helpful comments in their role as members of my advisory committee, and Robert O. Crowder and

Michael Davis for their helpful comments in their role t o an unconditioned Stimulus (US). Wagner

applicability of this reasoning to Slgnal-pro- duced variation in the immediate response

as readers on my committee

Requests for reprints should be sent to Nelson H.

Donegan, who is now at the Department of Psychology, Stanford University, Stanford, California 94305.

(1976 )

v

'

suggested t h at the response decre-

ee

r

ment during

uted to a transient priming in STM by recent

be attnb-

.

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habituation

, .,

,

,.

.,

may

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

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 maintainedwith 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).

BY CS PRIMING

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, requiringa graphicdeflection 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 isolationcham- ber for approximately 2 hr. During this period subjects' heads were shaved and the shock electrodes were im- planted. The experimentaldesigncalled 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 intervalsranging

from 90 to 150 sec (M

subjects, the auditory CS was reinforced (CS A + ), and

the vibratory CS was nonreinforced (CS B ~). For the remaining subjects, the vibratory CS was reinforced

(CS A

During each session five test trials were introduced on

which CS B 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

ITI = 120 sec). For half of the

+

), and the auditory CS was nonreinforced (CS B ~).

298

NELSON H. DONEGAN

during the course of discrimination training. In each session one test trial (CS B 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., CS B was reinforced and CS A was nonreinforced. As in Phase 1, five test trials, presentations of CS A followed by the US (written herein as CS A ~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 CS A ~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 CS B

training trials, which occurred in an ABBA sequence,

in each of eight sessions. During each session three kinds

of test

trials. Each subject received six presentations of CS A reinforced at a 1-sec CS—»US interval (time from CS onset to US onset) and six presentations each of CS A reinforced and CS B reinforced for which the CS—.US interval was .5, 2, 4, or 8 sec in different sessions. The CS A and CS B 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.

+

and 54 CS A "discrimination

trials were introduced, one after every sixtraining

Results and Discussion

Discrimination and reversal training. The effects of discrimination training on subjects' conditioned responding to CS A and CS B 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 CS A

) increased across sessions, and the

mean percentage CRs to CS B (CS~) initially increased and then decreased across sessions.

Only in the last block of sessions of this phase was the mean percentage CRs to CS A

(CS

+

greater than the mean percentage CRs to

CS B . In reversal training, the mean per-

centage CRs to CS A (now, CS~) decreased,

and the mean percentage CRs to CS B (now, CS + ) increased across sessions. In the first block of sessions, the mean percentage CRs to CS A was greater than to CS B , 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 CS B was greater than to CS A . Results of statistical analyses of the initial acquisition data indicated that there was a reliable interaction of trial type (CS A vs. CS B ) with sessions, F(l, 42) = 5.26, p < .01. Subsequent paired comparisons indicated that the percentage CRs to CS A was not re- liably different from that to CS B 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 CS A was reliably greater than the percentage CRs to CS B 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 CS A + US and 10 CS B "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 (CS A vs. CS B ) 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 CS A ~US and

US trials in the first block of two

sessions were 6.0 mm and 5.4 mm, respec-

= .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 (CS A vs. CS B ) with sessions was reliable, F(l, 42) = 7.42, /x.Ol.

Tests 1 and 2. The mean percentages

100 CS B

+

tively, t(l) = 2.25, p

UR MODIFICATION BY CS PRIMING

299

 

ACQUISITION

REVERSAL

100

 

•-•

CS A +

•-•

CS A -

o-o

CS B -

o-o

CS B +

80

60

40

20

123

4

123

4

BLOCKS

OF

2

SESSIONS

Figure 1. Mean percentage eye blink CRs in acquisition, during which CS A served as CS + and CS B

served as CS~, and in reversal training, during which CS B pairs of sessions in Experiment 1.

as CS + and CS A served as CS~, for

served

CRs to CS A and CS B 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 CS A served as CS + and CS B as CS", appear on the left-hand side of Figure 2. As can be seen, preceding the US by CS A 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 (CS A , CS B , and US alone) were reliable, F(2, 12) = 12.72, p < .01. The amplitude of re- sponding on CS A trials was reliably greater than on CS B or US-alone test trials,

t(l)

= 3.40, p < .05 and t(l) = 4.12, p <

.01,

respectively, and responding on CS B

trials was reliably greater than on US-alone

served

trials, t(l) = 2.75, p

< .05.

The results of Test 2, in which CS

B

as CS + and CS A served as CS~, appear on

the right-hand side of Figure 2. As can be seen, preceding the US by CS B facilitated responding relative to preceding the US by CS A , 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 (CS A , CS B , and US-alone) were reliable, F(2,12) = 43.12, p < .01. Subsequent paired comparisons indicated that the amplitude of responding on CS B trials was reliably greater than on CS A or US-alone test trials, t(7) = 4.57,;? < .01 and t(l) = 8.69,p < .01, respectively, and that responding on CS A 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

12.5

E

J 10.0

ui cc 13

o U)

_J o

2

7.5

5.0

2.5

NELSON H. DONEGAN

TEST

1

CS B -

NONE

CS A -

TEST 2

CS B +

CS PRECEDING THE US

NONE

Figure 2. Mean amplitude of the eye blink response at the time of US presentation when the US was preceded by CS 1 , 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-

IO.O

 

•— • US PRECEDED

BYCS +

O—O

US

PRECEDED

BYCS -

7.5

D—Q US ALONE

5.0

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-

tude at

-.75

-.50

-.25

SECONDS

0

FROM US

.25

ONSET

.50

l.O

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

301

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

•— • US PRECEDED BY CS -

0—0 US PRECEDED BY CS -

in

O

O

O

UJ

UJ

5

10 0

7 5

5.0

5

I 0

US

2.0

ALONE

4.0

8 0

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 respondingon CS + US

and CS~US trials averaged across the re-

maining, nontraining, intervals (.5, 2, 4, and 8 sec). The mean response amplitudesduring

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

US trials was reliably greater than on

trials at the 1-sec training interval,

CS

+

CS'US

302

NELSON H. DONEGAN

i io.o

or

g

3

o

D

^

Ul

UJ

1

UJ

5

7 5

5.0

2 5

•-•

CS +

o-o CS-

5

1.0

20

SECONDS

4.0

FROM CS

8 0

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

amplitude of the response at 8 sec after CS

onset. Figure 5 presents the average wave-

form of the CR on CS

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

and CS~ trials. As

the onset of CS +

and

CS"

+

as well

as

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

training interval. If groups of subjects were trained at .5-, 1-, and 2-sec CS—»US inter- 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- cally occurs, then each group should show 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.

+

, independently of the

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

303

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

diminished the amplitude

of the US by CS

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

+

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 malerabbits. Apparatus. The apparatus was modified so as to allow measurement of subjects' gross body movement. An Astatic Model 16 phonograph cartridgewas 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 restrainingbox. 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 +

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

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-

and the

1), where millimeterpen

+

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

two sessions of discrimination

testing, subjects received

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 approximatelyequated. 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 discriminationtrain- 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 trainingtrials, 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

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.

days of discrimination training.

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

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

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

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

and CS" in

. However, for lack

+

+

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

US discrimination training trials

and the 5 CS~US and 5 US-alone test trials

the 50 CS

+

z

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o

£

CL

<

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100

80

60

10

-

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.

BY CS PRIMING

£

E

Ul

CC.

3

O

O

S

22.5

20.0

17.5

15.0

CS+

CS-

NONE

CS

PRECEDING THE

US

305

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

the US in each of the trial types of Test 1 was determined as in Experiment 1. Figure 8 shows the average eye blink response am-

plitude at various points in time in relation

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

to US onset on CS

+

306

NELSON H. DONEGAN

20.0

17 5

•— • US PRECEDED BY CS +

0-0 US PRECEDED BY CS -

15.0 a—o US ALONE

12.5

10.0

7.5

5.0

2.5

0

-I.

-7 5

-.50

-.25

0

.25

50

l 0

 

SECONDS

FROM

US

ONSET

Figure 8. Amplitude of the eye blink response prior to, during, and after presentation of the US when

, 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 .05sec.)

the US was preceded by CS

+

immediately prior to the US onset was

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.

greater on CS

+

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

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o:

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•— • US PRECEDED

0—0 US

PRECEDED

BYCS +

BYCS -

 

US ALONE

UJ

5

 

2

CS +

CS-

CS

PRECEDING

NONE

THE

US

Figure 9. Mean amplitude of the movement response at the time of US presentation when the US was pre- ceded by CS + , CS~, or neither CS in Test 1 of Exper- iment 2.

cs-us

INTERVAL (sec)

Figure 10. Mean amplitude of the eye blink response at the time of US presentation when the US was pre- ceded by CS + or CS~ at CS^US onset intervalsranging 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.)

UR MODIFICATION BY CS PRIMING

307

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

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

mean amplitudes of responding on CS

+

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•— • US PRECEDED BY CS +

0—0 US PRECEDED 8Y CS -

. 5

1.0

US ALONE

2 0

CS-US

4. 0

INTERVAL(sec)

8. 0

Figure II . 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

5102 0

SECONDS

4.0

FROM CS

8.0

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

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.

+

and CS" as well as the amplitude of the

+

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

trials was greater than the

response on CS

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

US

INTENSITY

(mA)

6.0

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

309

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 intensitieswere 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'

UJ

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CO

UJ

K.

2.0

•— • US

PRECEDED BY CS ^

0—0 US

PRECEDED BY CS -

D—D US ALONE

US

INTENSITY (mA )

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.

movement responses during the US period

are shown in Figure 14. Preceding the US

by CS + acted

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-

to decrease the amplitude of

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

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

amplitude across test trial types (CS

+

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

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

t(7) = 5.78, p<.01 , respectively,

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-intensityUS 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

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 mimickingresponse 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

US trials is likely to be acombination

+

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 simplehuman 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-

STM.

That is to say, CS +

activation of the US representation and re- sults, for example, in eye blink and move- ment CRs that can be observed. Subsequent

presentation produces

ity of a unitary US representation in

UR MODIFICATION

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:

BY CS PRIMING

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

Search Opens for Editor of JEP: General

The Publications and Communications Board has opened nomi- nations for the editorship of the Journal of Experimental Psy- chology: General for the years 1984 through 1989. Candidates must be members of APA and should be available to start receiving manuscripts in early 1983 to prepare for issues published in 1984.

To nominate candidates, prepare a brief statement of one page or less in support of each nomination. Submit nominations no later than January 1, 1982, to the Chair of the Search Committee:

Janet T. Spence Department of Psychology University of Texas at Austin Austin, Texas 78712