Changes in the retrievability of associations to elements of the compound CS determine the expression of overshadowing

In two experiments, rats were presented with a taste conditioned stimulus (CS) alone, an odor CS alone, or an odor-taste compound followed by lithium chloride injection. When tested 1 day following conditioning, there was evidence that the odor cue overshadowed conditioning to the taste; however, there was no indication of overshadowing following a longer (21-day) retention interval, despite undiminished strength of the aversion in animals conditioned with only the single element (taste). The overshadowing observed at the 1-day retention interval was not reciprocal. Rats conditioned with the odor CS alone or with the compound CS expressed odor aversions of comparable strength—that is, no overshadowing. However, in contrast to the taste aversion, overshadowing of conditioning to the odor by taste was evident following a 21-day retention interval. Rather than reflecting a failure of the overshadowed stimulus to acquire associative strength, these data suggest that overshadowing may be expressed, or not expressed, as a result of changes in the relative retrievability of learned associations over time.     

Overshadowing of explicitly unpaired conditioned inhibition is disrupted by preexposure to the overshadowed inhibitor

In two conditioned lick-suppression experiments with rats, the interaction of preexposure to a target stimulus and subsequent overshadowing of conditioned inhibition treatment was examined. Blaisdell, Bristol, Gunther, and Miller (1998) have demonstrated that stimulus preexposure and overshadowing counteract each other in their effects on Pavlovian excitation, producing strong excitatory responding to the target stimulus. In the present experiments, a conditioned inhibition analogue of this effect was examined. Overshadowing of conditioned inhibition (i.e., attenuation of inhibitory control of behavior by the target stimulus) was demonstrated when a more salient stimulus was compounded with the target inhibitor during conditioned inhibition training. However, the overshadowing deficit was attenuated by preexposing the target stimulus prior to overshadowing treatment. To account for this phenomenon, we contrast the comparator hypothesis (Miller & Matzel, 1988) with contemporary models that focus on associative acquisition.     

Taste and odor in conditioned flavor preference learning

Almond and peppermint extracts were combined with salt and citric acid as cues in conditioned flavor preference conditioning. In Experiment 1, extracts overshadowed tastes, although tastes and extracts conditioned equally well when presented in isolation. In Experiments 2 and 3, tastes and extracts were conditioned in isolation prior to conditioning of a taste/extract compound. The conditioning history of the tastes and extracts did not affect the overshadowing of taste by extract. The results of Experiment 4 showed that rats could learn to discriminate between a taste and extract presented in isolation vs. the taste/extract compound. Thus, extracts do not interfere with sensing the tastes. We suggest that a taste/extract compound produces a configural stimulus that is more characteristic of the extract than the taste.     

Extinction of the overshadowing CS after overshadowing in conditioned taste aversion

Using a conditioned taste aversion preparation overshadowing of flavor-illness association was produced through the presentation of a second flavor during the interval between the first flavor and illness. The modulatory effects of extinguishing the association between the second (over-shadowing) flavor and illness on conditioned responding to the target flavor was investigated. In Experiment 1, we found that, following one-trial overshadowing, extinction of the overshadowing flavor had no effect on conditioned responding to the target flavor. In Experiment 2, we found a similar absence of an effect of extinction of the overshadowing stimulus in a multitrial over-shadowing paradigm. Experiment 3 confirmed the results of Experiments 1 and 2 using conditioning parameters that were designed to weaken the association between the overshadowed flavor and illness. In Experiments 4 and 5, we used simultaneous presentation of the flavors during conditioning and obtained a weakened aversion to the overshadowed flavor when the overshadowing CS was extinguished. These findings are inconsistent with previous observations in conditioned fear preparations that suggest that extinction of the association between the overshadowing stimulus and the unconditioned stimulus attenuates overshadowing. Possible reasons for the discrepant results are discussed.     

Covariation in conditioned response strength between stimuli trained in compound

Conditioned lick suppression by water-deprived rats was used to elaborate on recent evidence that the attenuated conditioned response elicited by an overshadowed stimulus may be enhanced by extinction of the overshadowing stimulus with which it had been trained in simultaneous compound. Using a modified serial stimulus arrangement in which a light coexisted with the last half of a tone that terminated with footshock, it was found in Experiment 1 that the tone overshadowed the light. Extinction of the tone-shock association resulted in a virtually complete recovery of the response to the overshadowed light. Using this serial overshadowing procedure, the possibility that the strength of a conditioned response to an element trained in compound covaries as a function of the strength of the response to the other element was tested in Experiment 2. Following overshadowing training similar to that of Experiment 1, independent reinforcement of the overshadowed light, that is, associative inflation, was found to have no deleterious effect on the response to the overshadowing tone. This suggests that the effects of postconditioning extinction and inflation of one element do not have symmetrical effects upon responding to the other element. The results of Experiment 2 were replicated in Experiment 3 using a simultaneous compound stimulus as opposed to the serial compound of the previous studies. These results are discussed in terms of various associative and cognitive models of learning and performance.     

Overshadowing as a function of trial number: Dynamics of first- and second-order comparator effects

In two conditioned lick suppression experiments with rats, we examined the permanence of the overshadowing effect as a function of the number of compound reinforced training trials. In Experiment 1, robust overshadowing was observed following 4 compound-US pairings but dissipated with 36 pairings. Overshadowing decreased because responding to the overshadowed stimulus increased, not because responding by the control group decreased. This dissipation was stimulus specific and not attributable to a response ceiling. Experiment 2 extended the generality of the effect to a sensory preconditioning design and further demonstrated that overshadowing lost through many compound-US pairings was restored by posttraining extinction of the training context. The results are explicable in terms of the extended comparator hypothesis (Denniston, Savastano, & Miller, 2001) under the assumption that the impacts of first- and second-order comparator processes grow differentially as a function of number of trials.     

Overshadowing by food of stimulus control by a keylight in a two-element serial compound

Pigeons were given successive discrimination training in which pecking during a choice period when the key was white was either reinforced or not, depending upon the prior presence or absence of a discriminative stimulus, which was a two-element serial compound. The compound consisted of a keylight and food, with food presented second or first in a forward or backward pairing for different groups of pigeons. In Experiment 1, the sequence was an S+ indicating reinforced trials, while in Experiment 2, the sequence was an S? indicating nonreinforced trials. Following acquisition of discriminated operant behavior, a sequence generalization test was administered during which all possible orders of the two stimuli were presented on test trials prior to the onset of the choice period. The results showed that food overshadowed stimulus control by the color of the light on the key on the sequence-generalization test, independently of whether food was presented first or second during training and independently of whether food was associated with reinforcement or nonreinforcement. The similarity of results for the two experiments suggests that overshadowing occurs independently of whether the compound is a discriminative stimulus for reinforcement or nonreinforcement. Simultaneous presentation of elements of a compound stimulus is not necessary for overshadowing because the phenomenon was captured with sequentially presented stimuli.     

Pavlovian appetitive discriminative conditioning inAplysia californica

Three experiments demonstrated Pavlovian appetitive discrimination learning in the marine mollusc,Aplysia californica. In each experiment, subjects were exposed to two conditioned stimuli; one stimulus (CS+) was paired with food presentations and the other stimulus (CS?) was never followed by food. In Experiments 1 and 3 different chemosensory stimuli were used, and in Experiment 2 different tactile stimuli were used. For both types of conditioned stimuli, bite responses occurred significantly more often to the CS+ than to the CS?. Experiment 2 also showed thatAplysia could learn a reversal of this discrimination. Experiment 3 showed that nonreinforced presentations of CS+ resulted in a decline in the frequency of conditioned biting. The implications of these results for neurobiological analyses of learning are discussed.     

Blocking in autoshaped lever-pressing procedures with rats

Rats will approach and contact a lever whose insertion into the chamber signals response-independent food delivery. This “autoshaping” or “sign-tracking” phenomenon has recently attracted considerable attention as a platform for studying individual differences in impulsivity, drug sensitization, and other traits associated with vulnerability to drug addiction. Here, we examined two basic stimulus selection phenomena—blocking and overshadowing—in the autoshaped lever pressing of rats. Blocking and overshadowing were decidedly asymmetrical. Previously reinforced lever-extension conditioned stimuli (CSs) completely blocked conditioning to auditory cues (Exps. 1 and 2), and previously nonreinforced lever-extension CSs overshadowed conditioning to auditory cues. By contrast, conditioning to lever-extension CSs was not blocked by either auditory (Exp. 3) or lever-insertion (Exp. 4) cues, and was not overshadowed by auditory cues. Conditioning to a lever-insertion cue was somewhat overshadowed by the presence of another lever, especially in terms of food cup behavior displayed after lever withdrawal. We discuss several frameworks in which the apparent immunity of autoshaped lever pressing to blocking might be understood. Given evidence that different brain systems are engaged when different kinds of cues are paired with food delivery, it is worth considering the possibility that interactions among them in learning and performance may follow different rules. In particular, it is intriguing to speculate that the roles of simple cue–reinforcer contiguity, as well as of individual and aggregate reinforcer prediction errors, may differ across stimulus classes.     

The overshadowing of instrumental conditioning by a stimulus that predicts reinforcement better than the response

In an experiment designed to demonstrate the overshadowing of appetitive instrumental conditioning, three groups of rats were given 10 sessions of RI (random interval) training in which reinforcement was delivered 450 msec following the response. The correlated group experienced a stimulus during the response-reinforcer delay interval, while an uncorrelated control group experienced a similar brief stimulus occasionally following responses, but these were not responses that typically produced reinforcement. A no-stimulus control group did not experience the brief response-produced stimulus. The experiment was run in two replications. The first employed a light as the critical stimulus, the second a tone. Over the 10 RI sessions, subjects in the two control conditions increased their rate of responding significantly faster than subjects in the correlated condition in both replications. This finding was interpreted as an instance of the overshadowing of the acquisition of signal value by a response because of the presence of the stimulus, which, in the correlated condition, was a more reliable predictor of reinforcement than the response. A subsequent conditioned reinforcement test confirmed that, in the correlated condition, the stimulus had, indeed, become a signal for reinforcement as a function of RI training.     

Overshadowing in the spatial domain

In Experiments 1 and 2, rats were trained in a Morris water maze to locate a hidden platform, the location of which in the circular pool was defined by four visual landmarks (A, B, C, and D), spaced at equal intervals around the edge of the pool. Control animals were trained with these four visual landmarks only. But for animals in the overshadowing groups, an auditory component, X, was added to Landmark D. Test trials, given at the end of training, consisted of placing the rat in the pool with no platform present and recording the time rats spent in the platform quadrant. In Experiment 1, the overshadowing group spent less time in the platform quadrant than controls when tested with D, but the two groups performed equally well on test trials that did not use D. We conclude that the auditory Component X overshadowed the visual Landmark D. In Experiment 2, we obtained evidence of reciprocal overshadowing, of D by X and of X by D. The results of Experiment 3 suggested that an appeal to generalization decrement might not be sufficient to explain these overshadowing effects.     

Conditioned reinforcement and complex discrimination: Can conditioned reinforcing value be specific to a complex of three cues?

The present experiment is concerned with the nature of the cues that might acquire conditioned reinforcing value, and the ways in which such cues might interact with one another. Red and green colored keylights were differentially paired with food dependent upon the houselight context (A or B) and the trial type (training or choice/forced). The duration of the colored keylights was varied between groups in an attempt to manipulate the effectiveness of the short-term memory of trial-type cues at the trial’s end. The red and green stimuli were of 30 sec duration for Group 30 and of 3 sec duration for Group 3. The results indicated that the choices of the pigeons in Group 30 were influenced by the houselight context present and by the keylight color. The choices of the pigeons in Group 3 seemed to be influenced by the houselight context present, the keylight color, and the memory of trial-type cues. Memory cues for trial antecedents were not overshadowed by presumably more salient external houselight stimuli for the pigeons in Group 3. Two alternative explanations for the results are discussed, and determined to be unlikely based on the results of an earlier experiment. The present results are related to a model of the conditioned reinforcing value of momentary stimuli and of transmission of conditioned reinforcing value.     

Biological significance attenuates overshadowing, relative validity, and degraded contingency effects

Miller and Matute (1996) showed that blocking is attenuated when the blocked conditioned stimulus (CS) is “biologically significant” (i.e., when the CS has the potential to elicit vigorous responding of any kind). To the extent that blocking is representative of cue competition, this finding suggests that biological significance protects CSs against cue competition effects in general. In the present experiments, we tested this possibility by examining the influence of biological significance of CSs on other examples of cue competition, namely, overshadowing, the relative stimulus validity effect, and the degraded contingency effect in rats. In Experiment 1, we found that intense auditory stimuli induced transient unconditioned lick suppression, thereby indicating that intense sounds were of high inherent biological significance. In Experiment 2A, we found that cues with high inherent biological significance were protected from overshadowing. In Experiment 2B, this finding was extended to cues with high acquired biological significance, which was obtained through prior pairings with a reinforcer of the valence opposite to that used in the overshadowing treatment. In Experiments 3 and 4, we found that cues with high inherent biological significance attenuated the relative validity effect and the degraded contingency effect, respectively. These results lend support to the view that biological significance (inherent and acquired) protects stimuli from cue competition effects, a finding that is problematic for many contemporary theories of learning.     

Overshadowing of geometric cues by a beacon in a spatial navigation task

In three experiments, we examined whether overshadowing of geometric cues by a discrete landmark (beacon) is due to the relative saliences of the cues. Using a virtual water maze task, human participants were required to locate a platform marked by a beacon in a distinctively shaped pool. In Experiment 1, the beacon overshadowed geometric cues in a trapezium, but not in an isosceles triangle. The longer escape latencies during acquisition in the trapezium control group with no beacon suggest that the geometric cues in the trapezium were less salient than those in the triangle. In Experiment 2, we evaluated whether generalization decrement, caused by the removal of the beacon at test, could account for overshadowing. An additional beacon was placed in an alternative corner. For the control groups, the beacons were identical; for the overshadow groups, they were visually unique. Overshadowing was again found in the trapezium. In Experiment 3, we tested whether the absence of overshadowing in the triangle was due to the geometric cues being more salient than the beacon. Following training, the beacon was relocated to a different corner. Participants approached the beacon rather than the trained platform corner, suggesting that the beacon was more salient. These results suggest that associative processes do not fully explain cue competition in the spatial domain.     

Competing sources of stimulus value in anticipatory contrast

In previous studies of anticipatory contrast, identical target components (A and B) preceded either a lower (extinction) or a richer schedule. Higher response rates occurred during the target preceding the lower rate of reinforcement, whereas preference was in favor of the target preceding the richer schedule. In Experiment 1, the response and preference measures were positively related when additional stimuli, with no reinforcement of their own, preceded the target components. The effect of these additional stimuli was presumed to be due to their overshadowing of the Pavlovian association between the target components and their following schedules. Experiment 2 also demonstrated a consistent relation between response rate and preference in a conditioned reinforcement procedure. In the absence of a strong Pavlovian association, anticipatory contrast, like other forms of contrast in free-operant procedures, reflects an increase in the value of the target component with an unchanged reinforcement schedule.     

Integration of cardiac responses to serial stimuli after Pavlovian conditioning in rats

The expression of cardiac responses to sequences of two sounds was studied in restrained rats following discriminative trace or delay conditioning. Stimuli paired with a tail shock 10 sec later (CS1) elicited conditioned bradycardia. Unpaired or neutral stimuli (CS0) elicited mostly tachycardia. Rats did not learn to suppress responding to nonreinforced sequences with an interval of 6 sec between sounds. Responses to the second stimulus were significantly augmented following a CS1 stimulus, but not following a CS0 stimulus. Real-time summation of simple responses provided a more complete and quantitative prediction of dual responses than did resetting or facilitation. These results extend the time range over which summation may be observed from less than 2 sec to at least 16 sec. They appear to be inconsistent with models involving competition between unitary representations of stimuli in short-term memory and suggest the existence of multiple stimulus traces with independent time courses.     

A comparison of learned aversions to gustatory and exteroceptive cues in rats

The experiment provides a direct comparison of the ability of subjects (rats) to associate gustatory and exteroceptive stimuli with illness. Previous experiments which have made similar comparisons between gustatory and exteroceptive cues have suffered from certain methodological problems involving stimulus control and compounding. The present experiment utilized a between-subjects design wherein half of the subjects had an auditory cue associated with poisoning and half had a taste cue. In both cases, the other cue was present, but was not predictive of poisoning, The auditory cue, like the taste cue, occurred only during drinking. This comparison was made in both an immediate and a delayed poisoning situation. The experiment found that while subjects were able to quickly associate a taste cue with illness, they were unable to form a similar association between poisoning and the exteroceptive stimulus. Results also showed that subjects will fail to acquire a taste aversion to a novel and salient gustatory cue when that cue is followed by illness only 50% of the time. This latter effect was more pronounced in the delayed poisoning situation.     

Contrasting predictive and causal values of predictors and of causes

Three experiments examined human processing of stimuli as predictors and causes. In Experiments 1A and 1B, two serial events that both preceded a third were assessed as predictors and as causes of the third event. Instructions successfully provided scenarios in which one of the serial (target) stimuli was viewed as a strong predictor but as a weak cause of the third event. In Experiment 2, participants’ preexperimental knowledge was drawn upon in such a way that two simultaneous antecedent events were processed as predictors or causes, which strongly influenced the occurrence of overshadowing between the antecedent events. Although a tendency toward overshadowing was found between predictors, reliable overshadowing was observed only between causes, and then only when the test question was causal. Together with other evidence in the human learning literature, the present results suggest that predictive and causal learning obey similar laws, but there is a greater susceptibility to cue competition in causal than predictive attribution.     

Surprise and change: Variations in the strength of present and absent cues in causal learning

It is said that “absence makes the heart grow fonder.” But, when and why does an absent event become salient to the heart or to the brain? An absent event may become salient when its nonoccurrence is surprising. Van Hamme and Wasserman (1994) found that a nonpresented but expected stimulus can actually change its associative status—and in the opposite direction from a presented stimulus. Associative models like that of Rescorla and Wagner (1972) focus only on presented cues; so, they cannot explain this result. However, absent cues can be permitted to change their value by assigning different learning parameters to present and absent cues. Van Hamme and Wasserman revised the Rescorla-Wagner model so that the α parameter is positive for present cues, but negative for absent cues; now, changes in the associative strength of absent cues move in the opposite direction as presented ones. This revised Rescorla-Wagner model can thus explain such otherwise vexing empirical findings as backward blocking, recovery from overshadowing, and backward conditioned inhibition. Moreover, the revised model predicts new effects. For example, explicit information about the absence of nonpresented cues should increase their salience (that is, their negative α value should be larger), leading to stronger associative changes than when no explicit mention is made of cue absence. Support for this prediction is detailed in a new causal judgment experiment in which participants rated the effectiveness of different foods’ triggering a patient’s allergic reaction. Overall, these and other findings encourage us to view human causal learning from an associative perspective.     

Contextual modulation of attention in human category learning

In a category-learning experiment, we assessed whether participants were able to selectively attend to different components of a compound stimulus in two distinct contexts. The participants were presented with stimulus compounds for which they had to learn categorical labels. Each compound comprised one feature from each of two dimensions, and on different trials the compound was presented in two contexts, X and Y. In Context X, Dimension A was relevant to the solution of the categorization task and Dimension B was irrelevant, whereas in Context Y, Dimension A was irrelevant and Dimension B was relevant. The results of transfer tests to novel stimuli suggested that people learned to attend selectively to Dimension A in Context X and Dimension B in Context Y. These findings contribute to the growing body of evidence that people can learn to selectively attend to particular dimensions of stimuli dependent on the context in which the stimuli are presented. Furthermore, the findings demonstrate that context-dependent changes in attention transfer to other categorization tasks involving novel stimuli.