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Specificity of Target Versus Feature
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Journal of Experimental Psychology: Copyright 1994 by the American Psychological Association. Inc.
Animal Behavior Processes 0097-7403/941$3.00
1994. Vol. 20. No. 1.51-65

Context-Specificity of Target Versus Feature Inhibition in a

Feature-Negative Discrimination

Mark E. Bouton and James B. Nelson

Four experiments with rats examined the effects of a context switch on inhibition that was acquired
during a feature-negative discrimination. A target conditioned stimulus was paired with food when
it was presented alone but occurred without food when it was combined with a feature stimulus.
A context switch following training did not disrupt inhibition conditioned to the feature. However,
responding to the target was more difficult to inhibit when it was tested in a different context. It
is suggested that both the target and the feature acquired inhibition and that the target's inhibition
was especially sensitive to the context. The feature may inhibit responding to the target (a) by
directly suppressing the representation of the food and (b) by activating the target's own inhibitory
association with food, which is at least partly context-specific. Implications for theories of inhi-
bition and negative occasion-setting are discussed.

The results of a number of experiments suggest that ex-
tinction performance is relatively specific to the context in
which it is learned. For example, if conditioned stimulus
(CS)-unconditioned stimulus (US) pairings are given in one
context and then CS-alone training (extinction) is given in
another context, a return of the CS to the original condi-
tioning context can renew extinguished performance to the
CS (e.g., Bouton & Bolles, 1979; Bouton & King, 1983;
Bouton & Peck, 1989). Although extinction performance is
specific to its context, conditioning performance is much less
so. When the context is switched after conditioning, there is
often relatively little diminution of the conditioned response
(CR) (e.g., Bouton & King, 1983; Bouton & Peck, 1989;
Bouton & Swartzentruber, 1986; Hall & Honey, 1989; Kaye
& Mackintosh, 1990; Kaye, Preston, Szabo, Druiff, &
Mackintosh, 1987).
It may be reasonable to suppose that the CS acquires a
second association during extinction and that this association
is especially sensitive to the context (Bouton, 1993). One
possibility is that the animal learns a new inhibitory asso-
ciation with the CS (e.g., CS-no US) in extinction. Such
learning has been proposed by several conditioning models
(e.g., Konorski, 1967; Pearce, 1987; Pearce & Hall, 1980;
Wagner, 1981; see also Pavlov, 1927) that assume that non-
reinforcement of the CS in extinction will cause acquisition
of a second inhibitory association instead of the unlearning
of the original CS-US association (e.g., Rescorla & Wagner,
1972). According to these models, at the end of extinction,
the CS has both excitatory and inhibitory values. On this
Mark E. Bouton and James B. Nelson, Department of Psychol-
ogy, University of Vermont.
This research was supported by Grants BNS 89-08535 and mN
92-09454 from the National Science Foundation.
Correspondence concerning this article should be addressed
to Mark E. Bouton, Department of Psychology, University of
Vermont, Burlington, Vermont 05405.
51
view, the context-specificity of extinction could come about
if inhibition is relatively specific to the context in which it
is learned.
The present experiments were therefore designed to test
whether conditioned inhibition is reduced with a change of
context. To produce inhibition, we used a feature-negative
discrimination procedure. In this sort of procedure, a target
CS, such as a tone, is paired with a US when it is presented
alone (T +) but is presented without a US when it is combined
with a "feature" stimulus, such as a light (LT-). The feature-
negative procedure is widely known to produce inhibition to
the feature (L). Indeed, it is sometimes taken as the basic
means of producing conditioned inhibition (e.g., Rescorla &
Wagner, 1972). If conditioned inhibition is specific to its
context, then a context switch after feature-negative training
might cause a loss of inhibition to the feature CS.
Recent theories of conditioning, however, suggest that the
target stimulus might also acquire some inhibition in the
feature-negative discrimination. As is expected when a CS is
nonreinforced in extinction, the target itself might acquire an
inhibitory association when it is nonreinforced in compound
with the feature in this paradigm (e.g., Pearce & Hall, 1980;
Wagner, 1981). Thus, both feature and target may acquire
some inhibition when the compound is nonreinforced in the
feature-negative paradigm.
We designed the present experiments to investigate the
effects of switching the context after feature-negative dis-
crimination training. We used an appetitive conditiOning
preparation in which we monitored the number of times
the rat entered the food cup or magazine (magazine en-
tries) during the CS (e.g., Hall & Channell, 1985; Hall &
Honey, 1990; Kaye & Mackintosh, 1990; Wilson &
Pearce, 1990). The results suggest that inhibition is indeed
attenuated when the context is switched after feature-
negative training. However, in the present experiments, the
context switch reduced inhibition to the target CS but not
inhibition to the feature. The results have implications for
the issue of what is learned in feature-negative discrimina-
tion learning.
 52 MARK E. BOUTON AND JAMES B. NELSON
Experiment 1
The purpose of the first experiment was to assess some of
the properties acquired by the feature in the feature-negative
procedure to be used in Experiments 2-4. There were two
groups. Group FN (feature-negative training) received a
feature-negative procedure in which a tone was the target
(T+) and a houselight-off stimulus was the negative feature
(LT-). In this and all subsequent experiments, the elements
of the compound (L and T) were presented simultaneously.
In each session, an intermittent white noise was also rein-
forced (N + ). L was ultimately tested in compound with both
N and T at the end of training. If L had acquired true inhi-
bition, we expected it to inhibit responding to both T and N.
Notice that the transfer target (N) was never nonreinforced
before testing; if L inhibited responding to N, its inhibitory
effect could not depend on any inhibition conditioned to the
target.
A control group received the same T + and N + trials dur-
ing training, but L alone trials (L-) replaced LT-. This is a
somewhat conservative control for inhibition to L in Group
FN, because L- training could produce some inhibition to L.
However, consistent with several conditioning models (e.g.,
Rescorla & Wagner, 1972; Pearce & Hall, 1980; Wagner,
1981), more inhibition often accrues to a CS when it is
trained as a feature in the feature-negative arrangement (e.g.,
Rescorla & Holland, 1977).
Method
Subjects
The subjects were 16 male Wistar rats, bred at the University of
Vermont, that were approximately 110 days old at the start of the
experiment. They were individually housed in standard stainless
steel cages in a room maintained on a 18:6 hr Jight:dark cycle; the
experiment was conducted on consecutive days during the light part
of the cycle. The rats were food-deprived to 80% of their original
body weight throughout the experiment.
Apparatus
There were two counterbalanced sets of four conditioning boxes;
these provided two "contexts" in Experiments 2-4 but were not used
in that capacity here. Each box was housed in a sound attenuation
chamber and was constructed of clear acrylic plastic. The boxes
were 23 X 13 X 11 cm. The front and right side walls were trans-
parent, and the exterior of the back wall and left side were covered
with black paper. Illumination was provided by two 7.5-W white
incandescent bulbs mounted on the ceiling of the sound attenuation
chambers, 25 cm above the box floors. A small 1.2-W keylight (3
cm in diameter) was positioned behind the wall in the upper right
comer of the boxes, 12 cm above the floor and 3.5 cm from the right
wall. The rats were placed in the boxes through the ceiling.
On the right wall of each box a stainless steel recessed food cup
was positioned 3 cm above the floor and 1.5 cm from the back wall.
The food cup was accessed through a 6-cm square opening, and an
infrared photocell was mounted 3 mm behind the opening, 2.5 cm
from the bottom. In all boxes, inactive operant levers were located
on the back wall near the right comer 5 em above the floor. Back-
ground noise in all boxes was a constant 65 dB (re 20 /-LN/m2 [AD.
In one set of four boxes, the floor consisted of 3-mm bars
mounted parallel to the side wall, staggered so that the odd-
numbered bars were mounted 6 mm above the even-numbered bars.
Odd-numbered bars were 1.6 cm apart (as were the even-numbered
bars). The black side and back walls were lined with l-cm horizontal
white stripes spaced 1 cm apart. A dish containing 10 ml of a 4%
McCormick coconut extract solution (McCormick, Hunt Valley,
Maryland) was positioned near the food cup but was outside the box
on the floor of the sound attenuation chamber. The floors of the
second set of four boxes consisted of3-mm bars spaced 1.8 cm apart
mounted diagonally with respect to the chamber walls. A similarly
positioned dish of 2% McCormick anise extract solution provided
the distinctive scent. In summary, the two sets of boxes differed in
visual (stripes vs. no stripes), tactile (staggered narrowly spaced
bars vs. wide level bars), olfactory (anise vs. coconut scent), and
spatial (they were housed in different rooms) respects.
Four CSs were available. Two auditory CSs were presented
through a speaker mounted on the ceiling of the sound attenuation
chambers, 25 cm above the floor of each box (the speakers for each
box were identical). One CS was an 80-dB 3000-Hz tone (T), the
other was an intermittent (6 pulses per second) 65-dB white noise
(N). One of two visual cues (L) was provided by the offset of the
house lights, which caused complete darkness. The other visual CS
was the illumination of the keylight (K). The US was always two
45-mg food pellets (traditional formula, P. J. Noyes, Lancaster,
New Hampshire) delivered 0.2 s apart.
Procedure
The experiment was run on consecutive days. Each rat was run
consistently in the same box (counterbalanced).
Magazine training. The rats were initially placed in the boxes
for two 30-min exposure sessions on each of 2 days. On the next
day, they received two sessions of hand shaping in which they were
trained to approach and eat from the food cup at the sound of the
feeder's activation. The first and second sessions were approxi-
mately 30 and 15 min long, respectively. Each rat received a total
of about 40 pellets.
Conditioning. The animals were then randomly assigned to
two groups (n = 8). On each of the next 2 days, both groups received
one 75-min session in which T and N were each paired with the US
on six trials. CSs were always 30 s in duration; US onset was con-
tiguous with CS offset. Trials were presented in a double alternating
fashion (T+ T+ N+ N+ ... ) with the intertrial interval (ITI)
averaging 345 s.
Inhibition training was then carried out over the next seven 75-
min sessions. For both groups, each session contained 4 T + and 4
N + trials distributed evenly through the session. There were also
12 nonreinforced trials involving L. For Group FN, these were com-
pound trials in which L and T were presented simultaneously (LT-).
For the control group, L was presented alone (L-). Trial sequences
were changed daily. On any day, T + , N +, and nonreinforced trials
occurred in the same trial positions for both groups. The ITI was
variable with a mean of 195 s.
Test. On the day following the conclusion of inhibition train-
ing, both groups received a series of test trials designed to assess
L's effect on responding to T and N. Half the rats in each group
received three repetitions of the sequence LT-, T+, N+, LN-, L-,
and the other half received the sequence LN-, N+, T+, LT-, L-.
(The L-trials were included to assess possible second order con-
ditioning to L.) The test session lasted 56 min, and the average ITI
was 195 s.
Data collection and analysis. The computer recorded the num-
ber of magazine entries each rat made during the 30-s CS and during
 CONTEXT AND INHIBmON 53

the 3(}-s period immediately before the CS (the pre-CS period).
Responding during the CS was evaluated by analyses of variance
(ANOVAs), and planned comparisons were made using the methods
discussed by Howell (1987, pp. 431-443). Responding during the
pre-CS periods was analyzed with distribution-free tests because a
substantial number of scores of zero often made the distributions
nonnormal. Throughout, the rejection criterion was set at p < .05.
Results
Conditioning and inhibition training proceeded unevent-
fully. During testing, Group FN showed signs of conditioned
inhibition to L, whereas the control group did not.
Conditioning. Magazine entries during the conditioning
sessions are summarized in Figure 1. The groups were com-
pared on responding to the reinforced and nonreinforced
stimuli in separate ANOVAs. For the reinforced CSs, a Group
X CS (T or N) X Session analysis on the two initial con-
ditioning days that preceded actual inhibitory conditioning
revealed a main effect of session, F(1, 14) = 7.98; some
excitation was acquired before inhibition training began. All
effects and interactions involving group and CS were non-
reliable, Fs < 1. For the 7 days that involved inhibition trials,
a Group X CS (T or N) X Session ANOVA revealed no main
effects, Fs(l, 14) < 3.0, but there were significant Group X
Session andCS X Session interactions, Fs(6, 84) > 2.71. The
Group X Session interaction was the result of a reliable in-
crease in responding over sessions in Group FN, F(6, 84) =
3.89, with no such effect occurring in the control group, F
< 1. We do not have an explanation of this difference. The
CS X Session interaction took the form of an increase in
responding to N over training, F(6, 84) = 5.90, and respond-
ing to T did not increase reliably above that observed during
the two initial conditioning days, F(6, 84) = 1.75.
A Group X Session ANOVA on the nonreinforced stimuli
(LT- in Group FN and L- in the control group) confirmed
a group effect, F(l, 14) = 126.17, a session effect, F(6, 84)
= 6.14, and a Group X Session interaction, F(6, 84) = 3.01.
The interaction confirms that Group FN's initial responding
to LT declined overtraining, F(6, 84) = 8.70, and responding
to L in the control group was always low and did not change
significantly over sessions, F < 1. These data are consistent
with the possibility that initial responding to T became in-
hibited by L over sessions in Group FN.
Pre-CS responding during the phase averaged 1.32 for
Group FN and 0.84 for the control group. The difference was
not reliable, U(8, 8) = 15.
Test. Responding during testing is summarized in Figure
2, which shows the groups' mean responding during the first
presentations of the tone and noise targets alone and in com-
pound with L. (First trials are reported and analyzed because
of our emphasis on the first test trial in subsequent experi-
ments; analyses of session means supported identical con-
clusions.) The effects of L on responding to T and N were
evaluated in separate Group X CS-type (target vs. com-
pound) ANOVAs. The ANOVA comparing trials with the
tone target revealed a CS-type effect that fell short of sig-
nificance, F(I, 14) = 3.56, p = .08, and a significant Group
X CS-type interaction, F(I, 14) = 6.05. The group effect was
not reliable, F < 1. Simple effect comparisons confrrmed that
there was reliably less responding to the LT compound than
T in Group FN, F(l, 14) = 9.45, but not in the control group,
F<1.
The analysis of responding to the noise target revealed a
near-significant Group X CS-type interaction, F(l, 14) =
4.31, p = .057. Planned comparisons revealed significantly
less responding to the LN compound than N in Group FN,
F(1, 14) = 7.16, but no difference in the control group, F <

12
tn
U 10
Z
8 FN
CJ
-
Z
C
Z
6 CTRL

0 4
a..
tn
W
a: 2 LT- (FN)
0 ...... 0 --0--0--0-_ __ 0
0 L- (CTRL)
0
0 2 3 4 5 6 7 8 9

SESSION
Figure 1. Mean responding to the conditioned stimuli (CSs) during the conditioning sessions of
Experiment 1. N = intermittent noise CS; T = tone CS; LT = compound composed of houselight-
off and tone; L = houselight-off alone; FN = feature-negative group; CTRL = control group.
54 MARK E. BOUTON AND JAMES B. NELSON
TONE TARGET
10
tn
o 8
z
-
C!J 6
-Z
Z
Q
4
oD.
tn
W 2
a:
FN CTRL
Figure 2. Mean responding to the conditioned stimuli (CSs) during the first test trials of Experi-
ment 1. Tests with the tone and noise targets are shown at the left and right, respectively. Vertical
lines indicate standard error of the mean. FN = feature-negative group; CTRL = control group; L
= houselight-off CS.
1. Neither the CS-type nor group effects were significant, Fs
< 2.92. The results strongly suggest that L inhibited respond-
ing to both T and N in Group FN, but not in the control group.
Pre-CS rates averaged 1.72 and 1.57 for Group FN and the
control group on the trials just reported. There was no dif-
ference between the groups, U(8, 8) = 24, and no differences
in the individual groups' responding on excitor and com-
pound trials, smallest T(5) = 0, ps > .10.
Finally, the groups did not differ in their responding to L
alone, F(l, 14) < 1. The mean first-trial response was 1.00
and 0.88 for Group FN and the control group, respectively.
There was no evidence of second-order conditioning to L.
Discussion
The results of this experiment establish that the negative
feature (L) acquires inhibition in our feature-negative con-
ditioning procedure (Group FN). Significantly, L's inhibi-
tory effect transferred to a new stimulus (N). The fact that
the control group's L had no such effect suggests that
Group FN's result was more than simple generalization
decrement. Also, as N had never been nonreinforced prior
to testing, L's inhibitory effect also does not appear to de-
pend on the target (N) having had a possible inhibitory as-
sociation with the US. This result is consistent with the
view that the inhibitor acts, at least in part, by suppressing
the target CS's activation of a memory-representation of
the US (e.g., Rescorla & Holland, 1977).
Experiment 2
The second experiment was designed to ask whether in-
hibition conditioned to the feature is attenuated by a change
of context. The design is illustrated in Table 1. All subjects
NOISE TARGET
E:I Target Alone
o L + Target Compound
FN CTRL
received feature-negative training with the tone target in each
of two contexts. In Context A, houselight-off (L) was the
negative feature (i.e., the rats received T+, LT- training); in
Context B, illumination of the keylight (K) was the negative
feature (the rats received T +, KT - training). The design thus
equated the two contexts on their association with reinforce-
ment, nonreinforcement, and T. During the test, one group
received LT- in Context A, the context in which it had been
trained. The other group received LT- in Context B. Al-
though T had been nonreinforced in B throughout training,
this was the first time the inhibitory feature (L) was presented
there. If feature inhibition is specific to its context, L should
suppress performance to T in Context A, but not in Context
B. The experiment was designed to focus on the context-
specificity of learning to the houselight-offset CS (L) be-
cause complete darkness should be perceived as the same
stimulus in the two different contexts.
Method
Subjects and Apparatus
The subjects were 16 male Wistar rats, from the same stock as
in Experiment 1, which were 110 days old at the start of the ex-
periment. The apparatus was the same as in Experiment 1, but the
two sets of boxes now served as different contexts (counter-
balanced).
Procedure
The experiment was run on consecutive days. With the exception
of the final test day, there was one session in each context on each
day. Intersession interval was 75-85 min. For each rat, Context A
was provided by the context in which the first conditioning treat-
ment of the experiment was received.
 CONTEXT AND INHmmON
Table I
Designs of Experiments 2, 3, and 4
Group Conditioning
Experiment 2
A A (T+, LT-), B (T+, KT-)
B A (T+, LT-), B (T+, KT-)
Experiment 3
A A (L+, TL-), B (K+, TK-)
B A (L+, TL-), B (K+, TK-)
Experiment 4
A (T+, N+, LT-), B (T+, N+, KN-)
Note. A and B are contexts; T = tone conditioned stimulus (CS); L = houselight-off CS; K
key light CS; N = intermittent noise CS; + = reinforced trials; - = nonreinforced trials.
Magazine training. On the first day, all rats received two 30-
min sessions of box exposure (one per context) with the food cups
baited with food pellets to facilitate magazine training. On the next
day, each rat received a 3D-min session of magazine training (fol-
lowing the method used previously) in each context. Approximately
30 food pellets were delivered in each context.
Conditioning. On the next 2 days, the rats received excitatory
conditioning of T in both contexts. Both sessions on the first day
began with two nonreinforced exposures to the CSs to be used in
that context (T and L in Context A and T and K in Context B). The
ITI on these trials was 30 s. The next 12 trials were T+ trials, and
as usual, T terminated in US delivery. The IT! was variable with a
mean of 310 s; the sessions lasted 72 min. On the second day, there
were 12 more T+ trials in each context.
Inhibitory conditioning was then carried out over the next 9 days.
On each day there was a session of T +, LT- training in Context A
and a session of T + , KT- training in Context B. Each session was
60 min long and contained 4 T+ trials and 12 nonreinforced com-
pound trials (average ITI = 195 s). Trial positions of the reinforced
and nonreinforced trials were changed daily. On the first and third
day, all rats were run in Context A then in B; on the second day they
were run in B then A. On Days 4-9, half the rats received the A-B
sequence and the other half received B-A, with the first session for
each rat alternating daily between A and B. By the end of the phase,
the rats had had nine sessions in which L had been a negative feature
in A and nine sessions in which K had been a negative feature
in B.
Test. The rats were divided into two groups (n = 8) matched
on performance to T and L during inhibitory conditioning. (Box and
A-B sequence during training were also balanced in the new
groups.) A single test session was then conducted on the day fol-
lowing the last day of inhibitory conditioning. One group was tested
in Context A, and the other was tested in Context B. Both groups
received four repetitions of the trial sequence LT-, LT-, T +, T +.
The test was 60 min long and used the same IT! schedule that was
used in training. If feature inhibition is context-specific, we ex-
pected increased responding to LT in Context B.
Results
The rats learned the feature-negative discriminations in
Contexts A and B. However, when the L feature was tested
in a new context, there was no evidence that its ability to
inhibit performance to T was reduced.
55
Testing
A (LT-, T+)
B (LT-, T+)
A (TL-, L+)
B (TL-, L+)
A & B (T+, N+, LN-, KT-)
=
Conditioning. Figure 3 presents responding to T and the
inhibitory compounds during conditioning. The 9 days that
involved nonreinforced compounds were analyzed with a
Group X CS type (T vs. the inhibitory compounds) X
Context (containing either LT- or KT-) X Session
ANOVA. There was a main effect of CS type, F(l, 14) =
234.07, and a CS type X Session interaction, F(8, 112) =
15.71. Simple effect tests indicated that responding to T in-
creased over sessions, F(8, 112) = 11.97, and responding
to the compounds decreased, F(8, 112) = 7.30. No other
main effects or interactions were significant, Fs < 1.97, ps
> .097. The animals came to discriminate reinforced and
nonreinforced trials over training, the two discriminations
(T+, LT- and T+, KT-) were learned about equally rap-
idly, and performance did not differ between the groups.
Response rates during the pre-CS periods were 1.59 for the
group eventually tested in A and 1.65 for the group even-
tually tested in B, U(8, 8) = 34. Neither group had a dif-
ference between pre-CS responding on reinforced and non-
reinforced trials, Ts(8) > 10.
Test. Figure 4 summarizes performance to LT and T on
their first test presentations for the groups tested in Con-
text A and Context B. (We focused on the first test with
each stimulus because previous work with extinction pre-
dicted an effect primarily on the first trial.) L inhibited per-
formance to T in both contexts. A Group (context) X CS
type (LT vs. T) ANOVA revealed a strong effect of CS
type, F(l, 14) = 399.03; overall, there was less responding
to LT than T. Neither the group effect nor the Group X CS
type interaction approached reliability, Fs(1, 14) < 1.
Planned comparisons indicated no difference in responding
to the compound in the two contexts, Fs < 1, and no dif-
ference in responding to the tone alone, F(1, 26) = 1.32.
L's ability to inhibit T was not reduced when it was
switched to Context B. The animals tested in A and B had
pre-CS rates of 2.37 and 3.63, respectively, on the com-
pound trial, U(8, 8) = 31, and 0.88 and 2.62 on the
T-alone trial. The latter difference was significant, U(8, 8)
= 12, but as responding to T in the two contexts was en-
tirely consistent with the performance we had observed
 56 MARK E. BOUTON AND JAMES B. NELSON

10

en
o
8
-
z
e"
• T+ INA
-
Z
C
6
-- ....... - T+ IN B
Z ｾ＠ LT-INA
oQ. --<>-- KT-IN B
4
en
W
a:
o 2 3 4 5 6 7 8 9 10 11

SESSION
Figure 3. Mean responding to the conditioned stimuli (CSs) during the conditioning sessions of
Experiment 2. A = Context A; B = Context B; T+ = reinforced trials with the tone; LT- =
nonreinforced trials with the houselight-off/tone compound; KT- = nonreinforced trials with the
keylightltone compound.

throughout the experiment, there seems to be little need to
reinterpret CS responding. The groups did not differ in re-
sponding to either T or LT during testing.
Discussion
Whether L was tested in its usual context (A) or in a new
context (B), it strongly inhibited performance to T. The re-
sults provide no evidence that inhibition to the feature was
attenuated by a context switch in this procedure. With the
present method, inhibition to the feature CS transferred well
between the contexts.
reinforcement. However, in this experiment the animals re-
ceived L + ,LT- training in Context A and K + ,KT- training
in Context B. T was now a feature common to both contexts,
whereas Land K were targets uniquely connected with Con-
texts A and B, respectively. During testing, the rats received
LT and L in either Context A, the home context, or in Context
B. As in Experiment 2, our focus was the effects of the con-
text switch on the houselight-off CS (L), because complete
darkness should be perceived as the same stimulus in both
contexts. But in the present experiment L played the role of
the target, rather than the feature, in the discrimination that
was learned in Context A. If inhibition to the target is lost
with a context switch, we should observe an increase in re-
sponding to LT in Context B.

Experiment 3
We began the present experiments with the idea that in-
hibition learned to a CS during extinction might be specific
to the context in which it is learned. Although we began by
asking whether feature inhibition is context-specific (Experi-
ment 2), an extinguished CS may have more in common with
the target CS, rather than the feature CS, in a feature-negative
discrimination. Unlike a purely inhibitory feature, targets and
extinguished CSs share a mixed history of reinforcement and
nonreinforcement; both may have mixed excitation and in-
hibition (e.g., Pearce & Hall, 1980; Wagner, 1981). The third
experiment was therefore designed to ask whether inhibition
to the target, rather than to the feature, is attenuated when the
context is changed following feature-negative training.
The design, illustrated in Table 1, was the mirror image of
Experiment 2. During initial conditioning, Contexts A and B
were again home to feature-negative discriminations, and
were therefore associated with both reinforcement and non-
Method
Subjects and Apparatus
The subjects were 16 male Wistar rats from the same stock as in
the preceding experiments; they were approximately 120 days old
at the start of the experiment. Housing, maintenance conditions, and
apparatus were the same as in the previous experiments.
Procedure
The experiment was run on consecutive days following the two-
session-a-day procedure used in Experiment 2.
Magazine training. Box exposure and magazine training were
conducted following the procedure used in Experiment 2.
Conditioning. There were then 2 days in which L and K were
paired with the US in Contexts A and B, respectively. The two
sessions given on the first day each began with two nonreinforced
 CONTEXT AND INHIBITION
CONTEXT
12
U)
0 10
Z
8
CJ
-CZ 6
Z
0 4
D.
U)
W
a: 2
0
LT T LT
Figure 4. Mean responding to the conditioned stimuli (CSs) during the first test trials of Experi-
ment 2. Vertical lines indicate standard error of the mean. LT = houselight-off/tone compound; T
= tone CS.
exposures to the CSs to be used in that context (L and T in Context
A and K and T in Context B). The remainder of each session then
contained 12 trials in which the appropriate excitor (L in A and K
in B) was reinforced. On the second day, all rats received another
12 L + trials in A and 12 K + trials in B. Trial spacing was the same
as in Experiment 2.
Inhibitory conditioning was then conducted over the next 9 days.
On each day there was a session of L+, TL- in A and a session of
K+, TK- in B. The position of reinforced and nonreinforced trials
were changed daily and followed the schedule used in Experiment
2. The sequence of exposure to Contexts A and B was also organized
and balanced following the method used in Experiment 2.
Test. Rats were divided into two groups (n = 8) matched on
performance to Land TL over training. Once again, box and A-B
sequence given during conditioning were balanced over groups. On
the day following the end of inhibitory conditioning, both groups
received tests of Land TL. One group was tested in Context A; the
other group was tested in Context B. There were four repetitions of
the trial sequence TL-, TL-, L +, L +. Trial spacing was the same
as in Experiment 2.
Results
The conditioning phase proceeded uneventfully. Most im-
portant, during testing, target inhibition was lost: The rats
responded more to TL in a new context (Context B) than in
its original context (Context A).
Conditioning. Responding to the excitors and inhibitory
compounds during conditioning is shown in Figure 5. The
data from Days 3-11 (those that involved both excitor and
compound trials) were subjected to a Group X CS type (ex-
citor vs. inhibitory compound) X Context (containing either
57
A CONTEXT B
T
Lor K) X Session ANOVA. The analysis revealed a main
effect of session, F(8, 112) = 1.96, CS type, F(1, 14) =
13.87, and a CS type X Session interaction, F(8, 112) = 4.04.
There was more responding to the excitors than to the com-
pounds, and this difference increased over training: Simple
effects analyzing the interaction revealed both an increase in
responding to the excitors, F(8, 112) = 4.24, and a decrease
in responding to the compounds, F(8, 112) = 2.06, over days.
No effects involving context (Le., L vs. K) or group ap-
proached significance, Fs < 1.26, ps > .20. As in Experiment
2, Land K had roughly equivalent effects. The groups even-
tually tested in A and B had mean pre-CS rates of 1.56 and
1.71, respectively. These did not differ, U(8, 8) = 28, and
there was no difference in pre-CS responding on reinforced
and nonreinforced trials in either group, Ts(8) > 12.
Test. Figure 6 shows responding to TL and L on their first
presentations for the groups tested in Contexts A and B. A
Group X CS type ANOVA revealed a significant Group X
CS type interaction, F(1, 14) = 13.51; the difference between
responding to the excitor alone and to the compound clearly
depended on the context. The CS type main effect was also
reliable, F(I, 14) = 12.22, although the main effect of group
was not, F(1, 14) = 1.85. Simple effects revealed no dif-
ference in responding to L in Contexts A and B, F(1, 20) <
1. However, there was more responding to TL in Context B
than in Context A, F(1, 20) = 8.07. When the context was
changed, there was a substantial increase in responding to the
TL compound. The groups tested in A and B had mean
pre-CS rates of 1.25 and 2.25 on the test of TL and 2.88 and
1.13 on the trial with Lalone. Pre-CS rates did not differ on
either trial, Us(8, 8) > 16.
 58 MARK E. BOUTON AND JAMES B. NELSON
U)
0
Z 5
C!J
-CZ
Z
0 3
D.
U)
W
a:
0 2 3 4 5 6
SESSION
Figure 5. Mean responding to the conditioned stimuli (CSs) during the conditioning session of
Experiment 3. A = Context A; B = Context B; L+ = reinforced trials with the houselight-off CS;
K + = reinforced trials with key light CS; TL- = nonreinforced trials with the tonelhouselight-off
compound; TK- = nonreinforced trials with the tone/keylight compound.
Discussion
The results of this experiment suggest that performance to
the TL compound increased when it was switched from Con-
text A to Context B. As the feature (T) was not new to that
context, the effect was evidently due to some effect of the
switch on the target (L). One possibility is that inhibition to
the target was reduced when the context was switched fol-
lowing feature-negative training. The reduction was evident
primarily when the target was presented in compound.
Excitation to L was not reduced demonstrably with a con-
text switch in this experiment. This result is consistent with
previous research in this laboratory and in others (e.g.,
Bouton & King, 1983; Bouton & Peck, 1989; Hall & Honey,
1989; Kaye et aI., 1987; Kaye & Mackintosh, 1990) in sug-
gesting that excitation is not readily reduced by a context
switch. It should be noted, however, that previous reports
with the magazine-entry preparation have actually produced
conflicting evidence on this point (Hall & Honey, 1989; Kaye
& Mackintosh, 1990).
Experiment 4
The results of Experiments 2 and 3 begin to suggest that
a context switch reduces inhibition to the target (Experiment
3) more than inhibition to the feature (Experiment 2). The
purpose of Experiment 4 was to compare the effects of a
context switch on the target and the feature within a single
experiment.
The design is illustrated in Table 1. We used a within-
subjects method. Each rat received pairings of T and N with
food in both Contexts A and B. In addition, each excitor was
nonreinforced in compound with either L or K in one of the
two contexts. In Context A, Twas nonreinforced in com-
• L+ IN A
-- ...... - K+ IN B
--0-
---0--
TL-INA
TK-IN B
7 8 9 10 11
pound with L; in Context B, N was nonreinforced with K.
The subsequent test was designed to pit the effect of a context
switch on target inhibition against the effect on feature in-
hibition. Each target was tested in the context in which it had
been inhibited (the home context) and in the context in which
it had not been inhibited (the away context). When tested
away, the target was tested with a feature that was in its
conditioning (i.e., home) context. When tested in its home
context, the target was compounded with a feature that was
outside its conditioning context (i.e., away). If target inhi-
bition is more sensitive than feature inhibition to the effects
of a context switch, then we would expect more responding
to a compound when the target was tested away, even though
it was compounded with a feature that was at home. In this
experiment, the compound tests were transfer tests in which
targets and features were tested in new combinations.
Method
Subjects and Apparatus
The subjects were 16 male Wistar rats, from the same stock as
before, that were approximately 90 days old at the start of the ex-
periment. Housing, maintenance conditions, and the apparatus were
the same as in the preceding experiments. In this experiment, white
semitransparent paper covered the key light CS.
Procedure
The experiment was run on consecutive days with two sessions
(one in each context) on each day.
Magazine training. All rats initially received 15 min of expo-
sure to both contexts with the food magazines baited with food
pellets. On the next day, there were two 30-min sessions (one in
each context) in which the rats were trained to retrieve pellets from
 CONTEXT AND INHIBITION
CONTEXT A
8
U)
(J
6
z
C!J
3: 4
Q
Z
oQ. 2
U)
W
ｾ＠
TL L TL
Figure 6. Mean responding to the conditioned stimuli (CSs) during the first test trials in Experi-
ment 3. Vertical lines indicate standard error of the mean. TL = tonelhouselight-off compound; L
= houselight-off CS.
the magazine at the sound of the feeder. Approximately 20 food
pellets were delivered in each context.
Conditioning. On the next day, all CSs were preexposed in both
contexts. The rats were run in Context A and then Context B. The
sessions were identical and were 46 min long; each session con-
tained two repetitions of the sequence K-, T-, L-, N-, with an
average ITI of 3.5 min.
The next 2 days involved conditioning of both excitors in both
contexts. In each of the two daily 75-min sessions, there were six
T+ and six N + trials (average ITI = 345 s) given in a double
alternating fashion. On Day 1, all rats were run in Contexts A and
B in the sequence AB, and on Day 2, all rats received BA.
Over the next 18 days, the rats received inhibitory conditioning
treatments in the two contexts. Each of the two-daily 75-min ses-
sions contained 4 T+ trials, 4 N+ trials, and 12 trials in which a
compound stimulus was nonreinforced. In Context A, the nonre-
inforced compound was LT, and in Context B, the compound was
KN. On odd days of the phase, the rats were run in the context
sequence AB; on even days they received BA. The trial sequence
was changed daily; the average ITI was 195 s.
Test. On the final 2 days of the experiment, all animals were
tested with T, N, LN, and KT in both contexts. Half of the subjects
(n = 8) received testing in Context A on Day I and Context B on
Day 2; the other half received the reverse sequence. Each rat re-
ceived one session each day. In each session, the rats received four
repetitions of one of four stimulus sequences: KT, T, LN, N; LN,
N, KT, T; T, KT, N, LN; or N, LN, T, KT. The excitors were re-
inforced, whereas the compounds were nonreinforced. Half the rats
received an inhibitory compound first, and the other half received
an excitor first. Assignment to stimulus sequence was balanced with
respect to context sequence. Each test session was 60 min long with
an average ITI of 195 s.
Results
Conditioning proceeded uneventfully; the animals learned
to respond to the excitors and refrain from responding to the
59
CONTEXT B
L
compounds. During testing, there was more responding to the
compounds that contained targets away from their inhibited
contexts, even though the inhibitory feature was at home.
Conditioning. The results of the conditioning phase are
presented in Figure 7. For simplicity, the figure collapses
over CS identity. Some data were lost from four sessions (5,
10, 16, and 19), and these sessions were excluded from
the figure and the statistical analysis. Responding to the ex-
citors and inhibitory compounds was analyzed in separate
ANOVAs. Responding to the excitors during the inhibitory
conditioning sessions shown in the figure was put through a
CS (T vs. N) X Session type (in which a given excitor was
or was not nonreinforced) X Day ANOVA. The analysis
revealed a main effect of day, F(13, 195) = 7.61. There was
no CS type effect, F < 1. The CS X Day interaction was
reliable, F(13, 195) = 2.62, although simple effect tests ana-
lyzing the interaction revealed significant increases in re-
sponding to both T and N over days, Fs(13, 195) > 12.99.
No other main effects or interactions were significant except
for the session-type main effect, F(I, 15) = 39.08; overall,
there was less responding to the targets when they were pre-
sented in sessions in which they were also being nonrein-
forced. In principle, this effect could have been cued by con-
text, because each excitor was nonreinforced in only one
context. However, when we isolated responding on only the
first trials of the sessions, we found no difference between
responding to the targets in the nonreinforced context (mean
= 10.03) and reinforced-only context (mean = 9.27), F(1,
15) < 1. Thus, previous nonreinforced trials within the ses-
sion, rather than an anticipation cued by context, appeared to
cause the weaker average responding in the sessions in which
the targets were nonreinforced.
 60 MARK E. BOUTON AND JAMES B. NELSON

14

Target Away
en 12
0 /
Z
- 10 .... -... .... ,.
CJ
-Z
C
Z
8 " ..
Target Home
'

6
0
Q.
0"0 ,
en
w
't)
, .0
a:
4
't,' "''0 "0' ,,0... '0"0-0Compounds
0-0
"'''0'
2
0 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16

DAY
Figure 7. Mean responding to the conditioned stimuli (eSs) during the conditioning phase of
Experiment 4. Target Away = context in which the target was always reinforced; Target Home =
context in which the target was nonreinforced in compound with a feature.

Responding to the feature-target compounds was sub-
jected to a CS type (KN vs. LT) X Day ANOVA. The analysis
revealed a reliable effect of day, F(13, 195) = 33.73. The CS
type effect was also reliable, F(I, 15) = 13.86; there was
more responding to KN than to LT. The interaction was not
significant, F < 1.
The animals averaged 1.92 responses during the pre-CS
periods of the conditioning phase. There were no differences
in pre-CS rates to the excitors when they were in sessions
containing nonreinforced trials or not, and there was no dif-
ference in responding prior to the two types of inhibitory
compound trials, Ts(l5) > 36.
Test. Responding to the targets and feature-target com-
pounds during testing is shown in Figure 8. For simplicity,
the figure collapses over test order, which had no systematic
effect on the results, and over the identities of the excitors and
inhibitory compounds. The data were analyzed with a within-
subjects ANOVA that included context (target at home or
away), trial-type (target alone or in compound), target-type
(T and KT vs. N and LN), and trials as the factors. There was
significantly less responding to the compounds than to the
targets alone, trial-type main effect, F(I, 15) = 88.21. How-
ever, as suggested by the figure, the difference between
excitor and compound depended on the context, as indicated
by a Context X Trial-type interaction, F(I, 15) = 11.35.
Simple effect tests indicated that there was more responding
to the compound when the target was away than when it
was at home, F(1, 15) = 37.98; responding to the targets
alone did not differ between the contexts, F(l, 15) = 1.95.
(A separate but identical analysis of the first-trial
data confirmed a difference between the compounds, F( 1,
15) = 4.67, but no difference between the excitors tested
alone, F < 1.) These data make it clear that target inhibi-
tion was attenuated with a change of context in this proce-
dure. There were also significant effects of target type and
a Target-type X Trial-type interaction, F(l, 15) > 23.87,
which were consistent with the conditioning phase in sug-
gesting weaker inhibition by K than by L. No other inter-
actions involving target-type approached reliability. The
overall analysis also confirmed reliable effects of Trial,
F(3, 45) = 3.16, and a Trial-type X Trial interaction, F(3,
45) = 9.40; over testing, target-alone responding in-
creased, F(3, 45) = 7.42, and compound responding de-
creased, F(3, 45) = 6.07. The reliable increase in respond-
ing to the targets suggests that they were not at a response
ceiling at the beginning of testing. No other main effects or
interactions approached significance.
Pre-CS responding averaged 2.16 during testing. There
were no differences in responding in the two contexts on
either compound or target-alone trials, Ts(15) > 20.
Discussion
During testing, there was more responding to the com-
pounds when the targets were presented in the context in
which they had never been inhibited; inhibition was lost
when the target was tested away. This pattern was evident
even though it meant that inhibition was stronger in the con-
texts in which the features had never been presented before.
Somewhat paradoxically, there was greater inhibition when
the features were tested in a new context. These results
strengthen the conclusion suggested by Experiments 2 and 3:
In the present preparation, the target is more likely than the
feature to lose inhibition when the context is switched fol-
lowing feature-negative training.
Two details of Experiment 4 are worth noting. First, unlike
Experiment 3, in the present design visual cues provided
the features and auditory cues provided the targets. The
loss of target inhibition found here suggests that the com-
 CONTEXT AND INHIBITION
patible result in Experiment 3 was not due to its unique
use of an auditory feature and a visual target. Second, un-
like Experiments 2 and 3, the two visual CSs (L and K)
were balanced in the present design. We found a consistent
pattern of results with these cues; during testing, there was
more responding to LN in Context A than Context B (see
Table 1) and more responding to KT in Context B than
Context A. Such results help eliminate any concern raised
by the fact that L and K were not balanced in the previous
experiments. The present findings are general across sev-
eral different stimulus compounds.
Finally, it should be noted that the inhibition tests used in
Experiment 4 were transfer tests in which the elements were
combined in novel combinations. Evidently, the loss of target
inhibition with a context switch is general to tests with both
familiar (Experiment 3) and novel (Experiment 4) feature-
target combinations.
General Discussion
The present experiments examined the effects of a context
switch on the performance controlled by both the feature and
the target after feature-negative discrimination training. The
main result can be summarized simply: Performance to the
target stimulus was more difficult to inhibit when the target
was tested in a context in which it had never been inhibited
before (Experiments 3 and 4). The simplest explanation may
be to assume that the target acquires some type of inhibition
when it is nonreinforced during discrimination training (e.g.,
Pearce & Hall, 1980; Wagner, 1981) and that this inhibition
is lost at least partly when the context is changed. It is im-
portant to note that the loss of target inhibition was not ap-
12
tJ)
10
0
Z 8
CJ
-
Z
C
6
Z
0 4
Q. ｻ＾Ｍｾ＠ ....
tJ)
W 2
a:
0 +-__--r_ _---,r--_ _-r-_ _ｾｔ｡ｲｧ･ｴ＠
0 2
TEST TRIAL
Figure 8. Mean responding (::t:: one standard error of the mean) during the test trials of Experiment
4. Target Away = target in the context in which it had always been reinforced; Target Home = target
in the context in which it had been nonreinforced in compound with a feature; Feature Home =
feature in the context in which it had been trained; Feature Away = feature in the context in which
it had not been trained.
61
parent when the target was simply presented (tested) alone.
When the context was switched, we obtained a loss of target
inhibition that was primarily restricted to responding to the
feature-target compounds.
In contrast to inhibition to the target, inhibition to the fea-
ture appeared to transfer readily between our contexts. In
Experiment 2, a negative feature presented in a new context
was effective at inhibiting responding to its usual target there.
In Experiment 4, a negative feature presented in a new con-
text was effective at inhibiting a transfer target there. In both
cases, the feature inhibited a target that had previously been
inhibited in the new context. We do not argue that feature
inhibition cannot be reduced by a context switch with other
conditioning or testing procedures. But with the methods
used in the present experiments, a loss of inhibition to the
feature was surprisingly difficult to detect. The results of
Experiment 4, in addition to Experiments 2 and 3, suggest
that any effect of a context switch on inhibition to the feature
must be weaker than its effect on inhibition to the target.
The primary result, then, is that the context switch ap-
peared to reduce the target's "inhibitability." One possibility
is that the animal learned that the target was sometimes in-
hibited and that this property was lost with a context switch.
The "inhibitability" idea is similar to Holland's (1989b,
1992) view that animals store a CS in a "conditional memory
system" when it has been the target in an occasion-setting
discrimination. When a target is stored in this system, it is
recognized as having a conditional value. However, it is not
clear that a conditional memory system would allow a tar-
get's inhibitability to be specific to its context; why should
membership in the system be revoked with a context change?
We propose, instead, that the present results may follow more
Target Away
/
Target Home
/ Target Away, Feature Home
...... ｾＭﾢ＠
3 " 4
Home, Feature Away
 62 MARK E. BOUTON AND JAMES B. NELSON
Figure 9. Model of extinction. Tone (T) conditioned stimulus
has two types of associations with the unconditioned stimulus
(US): an excitatory association (arrow) acquired during condition-
ing and an inhibitory association (blocked line) acquired during
extinction. Inhibitory association is gated, and requires simulta-
neous input from context (A) and tone for activation.
directly from how inhibition may be represented in a general
memory system.
The present results are compatible with previous work on
the context-specificity of extinction (e.g., Bouton, 1991), and
we can begin an account of them by first considering ex-
tinction. Figure 9 presents one possible representation of
what is learned when a target tone CS has been paired with
a US and then extinguished. When the tone is initially paired
with a US, a CS-US association is learned. Conditioned per-
formance would depend on the CS's ability to activate the
representation of the US (e.g., Rescorla, 1973; Wagner,
1981). When the tone is then nonreinforced in extinction, a
second, new inhibitory association is learned (e.g., Pearce &
Hall, 1980; Wagner, 1981). Activation (or retrieval) of the
inhibitory association would interfere with performance oth-
erwise evoked by the excitatory association (e.g., Bouton,
1993). In contrast to the excitatory association, however, ac-
tivation of the inhibitory association is highly dependent on
the presence of the context present during extinction (A). In
Figure 9, input from both T and A converges on an inter-
mediate "control element" (e.g., Estes, 1976) that in tum
activates the inhibitory association. The control element is
assumed to function as an AND gate; complete activation of
the inhibitory link thus depends on the joint presence of the
CS (tone) and the extinction context (A). This model predicts
that extinction performance to T will be specific to the ex-
tinction context and is consistent with a memory-retrieval
analysis of extinction (Bouton, 1991, 1993). An extinguished
CS is "inhibitable" in the sense that it has acquired an in-
hibitory link, as well as an excitatory link, with the repre-
sentation of the US.
The same associative structure may be embedded in what
is learned during feature-negative training. Figure 10
presents a possible representation of the content of feature-
negative learning. As before, the target (T) enters into both
excitatory and inhibitory associations with the US, and the
inhibitory association is also linked, by the control element,
to its context (A). The new inhibitory feature (L), however,
is encoded so that it may inhibit responding to T through
activation of one of either of two links. In the first link, L may
suppress activation of the US representation directly (e.g.,
Rescorla & Holland, 1977). This link is suggested by the
results of Experiment 1, where an inhibitory feature inhibited
performance to a new CS that presumably had no inhibitory
association because it had never been nonreinforced. Several
theoretical approaches to conditioning have accepted this as
the mechanism of action for conditioned inhibitory stimuli
(e.g., Pearce & Hall, 1980; Rescorla & Holland, 1977;
Wagner, 1981).
The feature's second link is the new mechanism suggested
by the results of Experiments 3 and 4. Here the feature in-
hibits responding to the target by activating the target's own
inhibitory association with the US. This link operates
through the control element, which still functions as an AND
gate. After feature-negative training, target inhibition there-
fore depends on the joint presence of the feature, the target,
and the context in which the target's inhibition was learned
(A). Notice that the feature's effect through Link 1 does not
depend on context; this would allow the feature's action to
transfer partly between contexts (Experiment 4). However,
the action of the inhibitor through the target's inhibitory as-
sociation will depend on the target being in a context in which
its own inhibition has been learned (Experiment 3). This is
the important result obtained in the present experiments.
Figure 10. Model of a feature-negative discrimination. Tone tar-
get (T) conditioned stimulus has both excitatory and inhibitory
associations with the unconditioned stimulus (US). Negative fea-
ture (L) inhibits responding to T through two mechanisms: (1) a
direct inhibitory association with the US and (2) an excitatory link
with T's own inhibitory association. As in Figure 9, T's inhibitory
association is gated, in this case requiring input from three stimuli
for activation: the context (A), T, and L.
 CONTEXT AND INHIBITION
The feature's effect through Link 2 may be related to the
hypothetical actions of negative occasion setters (e.g., Hol-
land, 1985). Such stimuli are sometimes thought to act by
suppressing the target's own excitatory association with the
US (e.g., Holland, 1985; see also Rescorla, 1991). One im-
plication of the present experiments is that negative occasion
setters may instead activate the target's inhibitory associa-
tion. Another implication is that both of the links shown in
Figure 10 may be present following a single feature-negative
discrimination procedure. It seems possible, however, that
some procedures could encourage the encoding of one type
of linkage over the other. For instance, serial presentation of
feature and target (e.g., Holland, 1984), or a relatively weak
or nonsalient feature (Holland, 1989a), may encourage the
feature's effect on the target's inhibitory association (Link 2).
However, the present data suggest that both mechanisms of
action may be present to some degree in a single feature-
negative discrimination.
One problem with the scheme represented in Figure 10 is
that it does not account for all the facts surrounding a nega-
tive feature's ability to transfer and control responding to new
targets. To the extent that a negative feature can suppress the
US representation directly (Link 1), then inhibition would
transfer to any excitor associated with the same US (e.g.,
Rescorla & Holland, 1977). But transfer of the inhibition
mediated by Link 2 would clearly be restricted. As Figure 10
suggests, the second link could not affect responding to an
excitor that has no inhibitory association with the US. The
fact that responding to targets that have never been nonre-
inforced are not affected by a negative occasion setters (Hol-
land & Lamarre, 1984; Lamarre & Holland, 1987; Rescorla,
1985) may be consistent with this possibility. However, a
negative occasion setter's ability to suppress performance
does transfer to excitors that have been in a negative occasion
setting relationship (Holland & Lamarre, 1984; Lamarre &
Holland, 1987). The scheme represented in Figure 10 does
not explain why transfer should ever occur with Link 2. The
problem is significant, because transfer of Link 2 inhibition
appears to have occurred in Experiment 4, where a feature
inhibited a separately trained target when testing occurred in
the context in which the target had been inhibited.
The facts of transfer thus demand that a target's inhibitory
association be activated by separately trained feature stimuli.
One possible extension of the scheme presented in Figure 10
is illustrated in Figure 11. The figure models Experiment 4:
Two negative features (L and K) have been trained in feature-
negative discriminations with separate targets (T and N, re-
spectively) in separate contexts (A and B, respectively). As
in Figure 10, the features operate by two mechanisms. In one,
they suppress the US representation directly, and in the other
they activate the target's own context-specific inhibitory as-
sociation with the US. The only new idea is that the features
now first activate an intermediate unit (X) that in tum ac-
tivates the target's inhibitory link. If the separate features
converge on an intermediate unit, complete transfer of a fea-
ture's inhibition to a new target would be possible. Further,
the model predicts that transfer will be most complete if the
transfer target is encoded, as shown, with an inhibitory as-
sociation resulting from feature-negative training (Holland &
63
Figure 11. Model of two feature-negative discriminations that
allows the features to inhibit responding to new targets in the
targets' home contexts (Experiment 4). A tone target (T) and light
feature (L) have been trained in Context A (A); a noise target (N)
and keylight feature (K) have been trained in Context B (B). (Solid
and dashed lines merely separate the two discriminations.) The
features (L and K) can inhibit responding to T and N through two
mechanisms: (a) direct inhibitory associations with the uncondi-
tioned stimulus (US) and (b) excitatory links with an intermediate
unit (X) that in tum excites the targets' inhibitory associations with
the unconditioned stimulus (US). Each target's inhibitory associa-
tion with the US is gated and requires input from the target, the
intermediate unit, and the context for activation.
Lamarre, 1984; Lamarre & Holland, 1987). The intermediate
unit may actually embody what Holland first set out to cap-
ture in his conditional memory system: The inhibitory as-
sociations of various targets are combined together by a unit
that effectively labels them as inhibitable. Like Holland's
system, the present one suggests that transfer will be most
complete when the target has been inhibited in its own dis-
crimination. However, unlike Holland's system, the present
scheme further requires that the transfer target be tested in
a context in which it has been inhibited before. That was the
important new result of Experiment 4.
Pearce (1987) proposed a generalization model that pro-
vides a fundamentally different account of many phenomena
of feature-negative discrimination learning. In this model,
the animal learns excitation or inhibition to the aggregation
of cues present on any conditioning trial; performance to any
new combination of CSs then depends on the amount of
excitation and inhibition that generalizes from other condi-
tioned stimuli. Contextual stimuli have no special status in
the theory. Despite its success with some aspects of feature-
negative discrimination learning, the Pearce model does not
predict the present findings. In fact, our simulations suggest
that it often predicts the opposite of the results we obtained.
Consider Experiment 2. In that design (Table 1), T +, LT-
training in Context A would cause the learning of strong
excitation to the compound AT and inhibition to the com-
pound ALT. The treatment in Context B (T +, KT-) would
yield excitation to BT and inhibition to BKT. Test perfor-
mance to LT in Contexts A and B (ALT and BLT, respec-
tively) would then depend on generalization from all of the
 64 MARK E. BOUTON AND JAMES B. NELSON
conditioned stimuli. The degree of generalization is deter-
mined by the similarity between the conditioned and test
stimuli. Because similarity in the model depends on the pro-
portion of elements common to the conditioned and test
stimuli, excitation learned to BT should generalize heavily
to performance to BLT-lOO% of BT is in BLT. Generali-
zation of inhibition from BKT is weaker, because only 67%
of BKT is in BLT. The prediction for ALT is different. Ex-
citation would again generalize heavily from AT to ALT, but
here it is completely offset by inhibition conditioned directly
to ALT. The model thus predicts greater responding to BLT
than ALT during testing, a result that was not obtained in
Experiment 2.1
Now consider the Pearce (1987) model's account of Ex-
periment 3. In that experiment, L +, TL- training in A would
cause strong excitation to AL and inhibition to ATL, whereas
K +, TK- training in B should cause strong excitation to BK
and inhibition to BTK. Once again, the test compounds were
ATL and BTL. Excitation from AL should generalize heavily
to ATL, but this would be offset by inhibition conditioned
directly to ATL. In contrast, neither of the excitatory com-
pounds (AL and BK) would generalize heavily enough to
BTL (50% of each is in BTL) to offset the inhibition gen-
eralizing there from ATL and BTK (67% of each is in BTL).
Consequently, Pearce's model does not predict the strong
excitatory responding to BTL that was observed in Experi-
ment 3. In fact, the model predicts stronger responding to
BTL in Experiment 2 than in Experiment 3, the opposite of
what we actually obtained. Simulations of Experiment 4 also
produced predictions that reversed the actual outcomes: The
model predicted strong excitation to the targets in the con-
texts in which they were also inhibited, and this excitation
was expected to generalize most heavily to (and cause stron-
gest responding to) the compounds containing targets in their
inhibitable contexts. In actuality, however, those compounds
elicited the weakest responding. The Pearce model does not
predict the results of Experiments 2-4.
To our knowledge, the present data are not anticipated by
any existing interpretation of feature-negative discrimina-
tions. We know of no theory that predicts that target in-
hibitability, but not feature inhibition, will be specific to its
context. The present experiments have produced new evi-
dence that both the feature and the target may acquire in-
hibitory associations with the US in the feature-negative
paradigm. Although the feature's ability to inhibit targets
appears to transfer across contexts, the target's inhibitability
does not. The results suggest that an inhibitory feature may
sometimes inhibit performance by activating the target's own
context-specific inhibitory association with the US.
1 We describe the percentage of elements in the training stimulus
that were common to the trained and tested stimuli. The Pearce
model also considers the percentage of elements in the test stimu-
lus that were shared; these were incorporated in our simulations,
but are left out of the discussion for simplicity.
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