Overshadowing and stimulus duration

In 3 experiments, the authors investigated the effect of stimulus duration on overshadowing. Experiments 1 and 2 examined responding to a target conditioned stimulus (CS1) when it was conditioned in compound with a coterminating overshadowing stimulus (CS2) that was longer, shorter, or of the same duration (the long, short, and matched groups, respectively). Equal overshadowing of conditioning to CS1 was obtained in all 3 conditions relative to a control group conditioned to the light alone. There were, however, differences in responding to CS2 as a function of its absolute duration. Experiment 3 examined the contribution of the food-food interval/CS onset-food interval ratio to these findings. In Experiments 1 and 2, the ratio differed for the overshadowing CS but not for the target CS. In Experiment 3, this arrangement was reversed, but the pattern of results remained the same. The implications of these findings for trial-based and real-time models of conditioning are discussed.     

Overshadowing of subsequent events and recovery thereafter

Four experiments using a conditioned lick suppression preparation with rats were conducted to examine whether overshadowing of subsequent events could be obtained in Pavlovian backward conditioning (i.e. unconditioned stimulus [US] before conditioned stimulus [CS]), and to determine whether such overshadowing could be reversed without further training with the overshadowed CS, as has been reported in overshadowing of antecedent events. In Experiment 1, a backward-conditioned CS overshadowed a second backward-conditioned CS. Two posttraining manipulations, extinction of the overshadowing CS (Experiment 2) and shifting of the temporal relationship of the overshadowing CS to the US (Experiment 3), increased responding to the overshadowed CS. These results constitute the first unambiguous demonstration of stimulus competition between subsequent events using first-order conditioning, and they show that, like overshadowing with forward conditioning, such overshadowing is due, at least in part if not completely, to a failure to express information that had been acquired.     

Stimulus and temporal cues in classical conditioning

In 2 experiments, separate groups of rats were given stimulus conditioning, temporal conditioning, untreated control and (in Experiment 2) learned irrelevance control procedures, followed by a compound with both stimulus and temporal cues. Stimulus conditioning consisted of a random 15-s duration conditioned stimulus (CS) followed by food; temporal conditioning consisted of food-food intervals of fixed 90 s (Experiment 1) or fixed 75 + random 15 s (Experiment 2). The stimulus group abruptly increased responding after CS onset, and the temporal group gradually increased responding over the food-food interval. When the food-food interval was fixed 90 s, the temporal cue exerted stronger control in the compound, whereas when the food-food interval was fixed 75 + random 15 s, the stimulus cue exerted stronger control. The strength of conditioning, temporal gradients of responding, and cue competition effects appear to reflect simultaneous timing of multiple intervals.     

Facilitation of extinction by an increase or a decrease in trial duration

Five experiments examined the effects of altering the duration of a conditioned stimulus (CS) for extinction. For the first 3 experiments, rats received conditioning with a 10-s CS before different groups received extinction with a CS that was either the same duration or longer than that used for conditioning. For the remaining 2 experiments, conditioning was conducted with a 60-s CS before different groups received extinction with a CS of either the same duration or a shorter duration than that used for conditioning. In all experiments, extinction progressed more readily when the CS duration was different for the 2 stages than when it was constant. The results are discussed in terms of rate expectancy theory and associative learning theory.     

Filial imprinting and associative learning

Evidence is reviewed that is relevant to an interpretation of filial imprinting as a form of classical or Pavlovian conditioning. For such an interpretation it has to be assumed that the imprinting object provides both the unconditioned stimulus (UCS) and the conditioned stimulus (CS). The difficulty for an experimental approach is that CS and UCS are always presented simultaneously and cannot be manipulated independently, making it virtually impossible to test phenomena such as latent inhibition and extinction. Existing evidence is consistent with a conditioning theory of imprinting, for instance with regard to the reversibility of filial preferences. Some hitherto unexplained findings can be accounted for in terms of overshadowing and blocking. Results of an experiment are discussed in which the occurrence of blocking was explicitly tested. The findings support an associative learning interpretation of imprinting, but further experiments are needed to be able to rule cut alternative explanations.     

The Influence of Retention Interval on the US Preexposure Effect: Changes in Contextual Blocking over Time

A number of studies manipulating the length of the interval between conditioning and testing indicate spontaneous recovery from overshadowing, suggesting that certain instances of overshadowing represent a deficit in memory retrieval rather than a failure of animals to form an association between the overshadowed stimulus and the US. The present series of experiments examined the influence of lengthening the retention interval on blocking, another stimulus selection phenomenon that is typically interpreted as an acquisition deficit. The results indicated that when subjects were tested shortly (3 days) after training conditioning to a taste blocked subsequent conditioning to an odor conditioned in compound with that taste (Experiment 1), whereas prior conditioning to an odor did not block subsequent conditioning to a taste conditioned in compound with that odor (Experiment 2). This pattern of results was essentially unchanged when testing occurred at a longer (21-day) retention interval. However, there was evidence of a US preexposure effect in Experiment 2 when subjects in the US ONLY control condition were tested at the 3-day retention interval, but not when testing occurred 21 days after conditioning. Experiments 3 and 4 examined whether this loss of the US preexposure effect over time might actually represent a change in the degree of contextual blocking as the retention interval is lengthened. Exposure to the conditioning context either during the interval between Phase 1 and Phase 2 of conditioning (Experiment 3) or prior to Phase 1 of conditioning (Experiment 4) alleviated this US preexposure effect suggesting that the loss of the US preexposure effect as the retention interval is lengthened observed in Experiment 2 is due to changes in the degree of blocking by contextual stimuli over time. The results are discussed in terms of differential susceptibility of forgetting of two functional roles played by a contextual stimuli in the current situation-context as a CS and context as a retrieval cue for other CS-US associations.     

Interaction between preexposure and overshadowing: Further analysis of the extended comparator hypothesis

Three experiments with rats used conditioned suppression of barpress to test predictions of the extended comparator hypothesis, which assumes that the effectiveness of (first-order) comparator stimuli in modulating responding to a target conditioned stimulus (CS) is itself modulated by other (second-order) comparator stimuli. Experiment 1 demonstrated that both pretraining exposure to the target CS alone (i.e., CS-preexposure effect, also known as latent inhibition) and pretraining exposure to a compound of the target CS and nontarget CS (i.e., compound-CS-preexposure effect) counteract overshadowing, and that posttraining deflation (i.e., extinction) of the overshadowing stimulus attenuates responding to the target CS when overshadowing is preceded by a CS-preexposure treatment (i.e., yields a CS-preexposure effect), but not when overshadowing is preceded by a compound-CS-preexposure treatment. Experiment 2 examined the consequences of posttraining associative inflation of the overshadowing stimulus or the preexposure companion stimulus following conjoint compound-CS-preexposure and overshadowing treatment. Experiment 3 examined the consequences of posttraining inflation of the overshadowing stimulus or the context following conjoint CS-alone preexposure and overshadowing treatment. The results support the expression-focused comparator view in contrast to recent acquisition-focused models of retrospective revaluation.     

Prevention of the degraded-contingency effect by signalling training trials

Presentation of unsignalled unconditioned stimuli (USs) interspersed among Pavlovian excitatory conditioning trials weakens conditioned responding to a target conditioned stimulus (CS; Rescorla, 1968). However, signalling these intertrial USs with another cue (a cover stimulus) has been shown to alleviate this degraded-contingency effect (e.g. Durlach, 1982, 1983). In contrast to signalling the inter-trial USs, the present experiments examined the effect on the degraded-contingency effect of signalling the target CS-US pairings. Experiment 1, using parameters selected to avoid overshadowing, found that consistently presenting a cover stimulus immediately prior to the target CS-US pairings during degraded-contingency training alleviated the degraded-contingency effect. Experiment 2 examined the underlying mechanism responsible for this cover-stimulus effect through posttraining associative inflation of the cover stimulus or the context, and found that inflation of the cover stimulus attenuated responding to the target CS (i.e. empirical retrospective revaluation). The results are discussed in terms of various acquisition- and expression-focused models of acquired responding.     

Associative changes with a random CS-US relationship

Three experiments examined conditioned magazine approach in rats when a positive unconditioned stimulus (US) bore a random relation to a conditioned stimulus (CS). Experiment 1 found that over the course of conditioning the CS initially elevated responding relative to the baseline but then lost the power to do so. Transfer tests revealed that a CS-US association developed early and persisted despite the decline in magazine responding. Experiment 2 confirmed the persistence of CS-US associations and found them to be more substantial when a different US occurred during the CS than in its absence. In Experiment 3, when the situation was exposed to US-alone presentations prior to introducing the CS, there was little evidence that a subsequent random relation between the CS and US produced an association between them. These results agree with those of blocking and overshadowing experiments using discrete CSs and support an interpretation of the random procedure in terms of competition between the background and CS for conditioning.     

Overshadowing and Blocking as Acquisition Deficits: No Recovery After Extinction of Overshadowing or Blocking Cues

The effects of extinction of the blocking or overshadowing stimulus on conditioned responding controlled by the blocked or overshadowed stimulus were examined in seven appetitive conditioning experiments with rats. The experiments differed in their designs, stimuli used, the amounts of conditioning and extinction training, and the levels of conditioned responding produced. In all cases, conditioned responding to the blocked or overshadowed cue was either unaffected or reduced by extinction of the blocking or overshadowing cue. These data are consistent with accounts of overshadowing and blocking that attribute those phenomena to acquisition deficits, rather than to retrieval failures.     

Counterconditioning of an overshadowed cue attenuates overshadowing

In 3 Pavlovian conditioned lick-suppression experiments, rats received overshadowing treatment with a footshock unconditioned stimulus such that Conditioned Stimulus (CS) A overshadowed CS X. Subjects that subsequently received CS X paired with an established signal for saccharin (CS B) exhibited less overshadowing of the X-footshock association than subjects that did not receive the X-B pairings (Experiment 1). Experiment 2 replicated this effect and controlled for some additional alternative accounts of the phenomenon. In Experiment 3, this recovery from overshadowing produced by counterconditioning CS X was attenuated if CS B was massively extinguished prior to counterconditioning. These results are more compatible with models of cue competition that emphasize differences in the expression of associations than those that emphasize differences in associative acquisition.     

Timing of fear expression in trace and delay conditioning measured by fear-potentiated startle in rats

In two experiments, the time course of the expression of fear in trace (hippocampus-dependent) versus delay (hippocampus-independent) conditioning was characterized with a high degree of temporal specificity using fear-potentiated startle. In experiment 1, groups of rats were given delay fear conditioning or trace fear conditioning with a 3- or 12-sec trace interval between conditioned stimulus (CS) offset and unconditioned stimulus (US) onset. During test, the delay group showed fear-potentiated startle in the presence of the CS but not after its offset, whereas the trace groups showed fear-potentiated startle both during the CS and after its offset. Experiment 2 compared the time course of fear expression after trace conditioning with the time course in two delay conditioning groups: one matched to the trace conditioning group with respect to CS duration, and the other with respect to ISI. In all groups, fear was expressed until the scheduled occurrence of the US and returned to baseline rapidly thereafter. Thus, in both trace and delay fear conditioning, ISI is a critical determinant of the time course of fear expression. These results are informative as to the possible role of neural structures, such as the hippocampus, in memory processes related to temporal information.     

Event representation in Pavlovian conditioning: image and action

In a typical Pavlovian conditioning experiment, a relatively insignificant event, the conditioned stimulus (CS), is paired with a biologically more meaningful event, the unconditioned stimulus (US). As a consequence of those pairings, the CS is thought to acquire response characteristics of the US. In this article I describe experiments with rats that suggest that under some circumstances, the CS acquires control of perceptual processing of the US, in the absence of that US itself. I present three kinds of evidence for this surrogate processing, which I liken to imagery or hallucination: (1) CSs come to control specific, sensory-evaluative responses normally evoked only by the USs; (2) CSs can substitute for USs in the establishment of new learning about those USs themselves; (3) CSs can substitute for USs in the modulation of conditioning to other events, either overshadowing (interfering) or potentiating learning, in the same manner as the USs themselves. Finally, I compare these data with evidence for conditioned sensation and imagery in humans, and suggest that imagery may be a very basic process, evolutionarily derived from perceptual and conditioning processes adapted to deal with remote or absent objects.     

Temporal specificity of extinction in autoshaping

Three experiments investigated the effects of varying the conditioned stimulus (CS) duration between training and extinction. Ring doves (Streptopelia risoria) were autoshaped on a fixed CS-unconditioned stimulus (US) interval and extinguished with CS presentations that were longer, shorter, or the same as the training duration. During a subsequent test session, the training CS duration was reintroduced. Results suggest that the cessation of responding during an extinction session is controlled by generalization of excitation between the training and extinction CSs and by the number of nonreinforced CS presentations. Transfer of extinction to the training CS is controlled by the similarity between the extinction and training CSs. Extinction learning is temporally specific.     

Blocking and nonsimultaneous compounds: Comparison of responding during compound conditioning and testing

Three blocking experiments were run using a conditional emotional response procedure with rats as subjects. In each experiment, following initial conditioning to a light stimulus, blocking rats were conditioned to a compound in which a short-duration (30-second) noise was superimposed on the terminal portion of the longer-duration light. In Experiment 1, the light was always three minutes in duration; in Experiment 2, the light was switched from a constant (three-minute) to a variable (0.5–5.0 minute) duration at the start of compound conditioning; in Experiment 3, the light was variable in duration throughout conditioning. Control rats received the same compound conditioning experiences as the blocking rats but did not receive prior conditioning to the light by itself. These temporal manipulations had strong and systematic influence on the rats’ pattern of responding during compound conditioning. Most notably, experience with a constantduration light resulted in both blocking and compound animals showing little conditional suppression to the early conditional stimulus portion of light by itself, followed by strong conditional suppression to the terminal 30 seconds of light plus noise. When testing was done with the noise in isolation subsequent to compound conditioning, however, an equally strong blocking effect was obtained across all the temporal manipulations. In all cases, the blocking animals showed much less conditional suppression to the noise than did the compound control animals. This lack of correspondence between the rats’ responding during compound conditioning and their response to the noise by itself is theoretically puzzling. The confounding effect of inhibition of delay may provide at least a partial explanation of this pattern of results.     

Anterograde amnesia for Pavlovian fear conditioning and the role of one-trial overshadowing: effects of preconditioning exposures to morphine in the rat

Four experiments studied anterograde deficits in Pavlovian fear conditioning following prolonged exposure to the mu-opioid receptor agonist morphine. Injections of morphine produced temporally graded anterograde amnesia characterized by deficits in contextual and conditioned-stimulus (CS) conditioning 1 or 7 days and selective impairment in CS conditioning 21 days after last injection. This anterograde deficit in conditioning did not recover across a retention interval, was absent when rats were tested immediately after conditioning, and required the presence of an auditory CS. These results suggest that anterograde deficits in Pavlovian fear conditioning emerged from differences in susceptibility to 1-trial overshadowing of context by CS.     

Odor-aversion learning in neonate rat pups: the role of duration of exposure to an odor

Five experiments employed 2- and 10-day-old rat pups to examine the ontogeny of odor-aversion learning. When duration of exposure to the to-be-conditioned odor stimulus (CS) was long, Experiments 1 and 2 demonstrated substantial and similar aversions across the two ages regardless of whether the toxin unconditioned stimulus (US) occurred simultaneously with, immediately before, immediately after, or some time after the CS. These experiments, therefore, did not support the claim of Rudy and Cheatle (Ontogeny of Learning and Memory. Hillsdale, N.J.: Erlbaum (1979) ) that the 2-day old pup suffers a deficit in its learning about the relation between successively presented events. The remaining experiments showed, however, that the 2-day-old pups did not acquire an aversion to a briefly presented CS, that the failure to condition this CS was not due to its overshadowing by contextual cues, and that the 10-day-old pup did acquire an aversion to that CS. The experiments have revealed, therefore, that there are age-related differences in the rat pup's ability to learn about the relation between a briefly-presented CS and the effects of a US.     

Conditioned stimulus informativeness governs conditioned stimulus-unconditioned stimulus associability

In a conditioning protocol, the onset of the conditioned stimulus ([CS]) provides information about when to expect reinforcement (unconditioned stimulus [US]). There are two sources of information from the CS in a delay conditioning paradigm in which the CS-US interval is fixed. The first depends on the informativeness, the degree to which CS onset reduces the average expected time to onset of the next US. The second depends only on how precisely a subject can represent a fixed-duration interval (the temporal Weber fraction). In three experiments with mice, we tested the differential impact of these two sources of information on rate of acquisition of conditioned responding (CS-US associability). In Experiment 1, we showed that associability (the inverse of trials to acquisition) increased in proportion to informativeness. In Experiment 2, we showed that fixing the duration of the US-US interval or the CS-US interval or both had no effect on associability. In Experiment 3, we equated the increase in information produced by varying the C/T ratio with the increase produced by fixing the duration of the CS-US interval. Associability increased with increased informativeness, but, as in Experiment 2, fixing the CS-US duration had no effect on associability. These results are consistent with the view that CS-US associability depends on the increased rate of reward signaled by CS onset. The results also provide further evidence that conditioned responding is temporally controlled when it emerges.     

Lithium-mediated disruptions of latent inhibition: Overshadowing by the unconditioned stimulus in flavor conditioning

In a series of experiments, the ability of a single lithium preexposure to disrupt CS effectiveness was assessed using a latent inhibition procedure. Lithium preexposure administered proximal (90 min) to a saccharin familiarization trial reduced latent inhibition whereas a similar administration more distal (360 min) to flavor familiarization failed to do so. Additional experiments demonstrated that this socalled “US overshadowing” effect was not attributable to sensitization (Experiment 2), excitatory backward conditioning (Experiment 3), or state dependency (Experiment 4). The implications of US overshadowing for proximal US-preexposure effects are discussed.     

Potentiation and overshadowing in preweanling and adult rats

Four experiments compared the aversion acquired by 18-day-old and 60-day-old rats to a flavor that was either tasted alone or in combination, simultaneously or successively, with another flavor when paired with illness. The purpose was to study temporal variables and theoretical issues pertinent to potentiation and overshadowing while investigating ontogenetic differences in these phenomena. When either the preweanlings or the adults were presented a simultaneous compound flavor (sucrose/coffee) followed by lithium chloride-induced illness, greater sucrose aversions were found than for animals conditioned on sucrose alone--that is, potentiation. Preweanlings demonstrated greater potentiation than did adults, whether the conditioned stimulus-unconditioned stimulus (CS-US) delay was 0 or 1 hr. This potentiation was eliminated by nonreinforced presentations of the alternative CS element. Potentiation was not seen when the two flavors were presented successively as the CS; instead, overshadowing occurred. Tests of configuring by extinction procedures indicated a tendency for these animals to form a configured representation of the simultaneous compound solution. This disposition for configuring tended to be more pronounced for preweanlings than for adults. Ontogenetic differences in response to configuration, CS saliency, and generalization decrement seem consistent with at least one model of potentiation and the ontogenetic differences in potentiation seen in the present experiments.