Conditioned suppression of a positively reinforced shuttle response

Rats were trained to run up and down an alleyway for sucrose reinforcement on a variable interval schedule. Differential aversive classical conditioning with auditory CSs was then conducted in a separate apparatus (“off the baseline”) prior to those CSs being presented while the subjects were responding for sucrose in the alleyway. Once the effects of the CSs had extinguished, shock was reintroduced following one CS but not the other (“on the baseline” differential aversive classical conditioning). Both “off the baseline” and “on the baseline” conditioning resulted in conditioned suppression to the CS followed by shock, but little effect of the CS followed by no shock was found. In the “on the baseline” phase, total suppression of baseline responding occurred at moderate US intensities, and this appeared to result from the subject avoiding the location at which he was last shocked. At lower values, both baseline response rate and relative suppression ratio were functions of US intensity. The results are discussed in relation to the effects found in similar experiments using avoidance baselines.     

The effects of signaling the US in backward conditioning: A shift from excitatory to inhibitory learning

The associative effects of “backward” US-CS pairings were compared when the pairing occasions were either consistently preceded by a well-trained CS+ or were unannounced. The investigation employed a conditioned emotional response procedure with rats, under conditions in which all subjects received the same schedule of shock USs, some signaled and others not, and the back-ward CS was arranged to follow either the former or the latter, in separate groups. The major finding was that although the backward CS became excitatory when it followed unsignaled USs, it became inhibitory when it followed signaled USs. This outcome, which is in line with prior findings of Wagner and Terry (1975), is in accordance with a “sometimes-opponent-process” model proposed by Wagner (1981). It is contrary to data reported by Fowler, Kleiman, and Lysle (1984) that may have resulted from a confounding of the different circumstances of backward conditioning with differences in the predictability of the US in the experimental context.     

Temporal encoding in trace conditioning

Conditioned lick suppression in rats was used to explore the role of timing in trace conditioning. In Experiment 1, two groups of rats were exposed to pairings of a CS (CS1) with a US, under conditions in which the interstimulus interval (ISI) that separated CS1 offset and US onset was either 0 or 5 sec. Two additional groups were also exposed to the same CS1→US pairings with either a 0 or a 5-sec ISI, and then received “backward” second-order conditioning in which CS1 was immediately followed by a novel CS2 (i.e., CS1→CS2). A trace conditioning deficit was observed in that the CS1 conditioned with the 5-sec gap supported less excitatory responding than the CS1 conditioned with the 0-sec gap. However, CS2 elicited more conditioned responding in the group trained with the 5-sec CS1-US gap than in the group trained with the 0-sec CS1-US gap. Thus, the CS1-US interval had inverse effects on first- and second-order conditioned responding. Experiment 2 was conducted as a sensory preconditioning analogue to Experiment 1. In Experiment 2, rats received the CS1?CS2 pairings prior to the CS1→US pairings (in which CS1 was again conditioned with either a 0 or a 5-sec ISI). Experiment 2 showed a dissociation between first- and second-order conditioned responding similar to that observed in Experiment 1. These outcomes are not compatible with the view that differences in responding to CSs conditioned with different ISIs are mediated exclusively by differences in associative value. The results are discussed in the framework of the temporal coding hypothesis, according to which temporal relationships between events are encoded in elementary associations.     

Reacquisition following extinction in appetitive conditioning

In four experiments utilizing an appetitive conditioning preparation, reacquisition of conditioned responding was found to occur both rapidly and slowly following extinction. In Experiment 1, acquisition of responding to a tone that had been conditioned and extinguished occurred more rapidly than acquisition in either a group that received equivalent exposure to the food unconditioned stimulus or a “rest” control group that received only exposure to the apparatus in the first two phases. However, reacquisition was impaired relative to acquisition in a “learning-experienced” group that had previously received conditioning and extinction with a different stimulus. Experiments 2 and 3 produced similar results, but also found that high responding during reacquisition was confined to trials that followed reinforced, rather than nonreinforced, trials. Experiment 4, in which very few initial conditioning trials were used, produced reacquisition that was slow compared with both learning-experienced and rest controls. The results are consistent with a role for sequential learning: Reacquisition is rapid when animals have learned that reinforced trials signal other reinforced trials.     

Transient variations in responding to Pavlovian conditioned stimuli have implications for the mechanisms of “priming”

Conditioned responding to a well-trained conditioned stimulus (CS) was assessed in an eyelid conditioning situation under three conditions of prior stimulation: when preceded by recent presentation of the same CS, when preceded by recent presentation of a different CS, and when preceded by no recent stimulation. As compared to the latter condition, prior presentation of the same CS depressed responding, while prior presentation of a different CS augmented responding. Both effects diminished as the interval separating the test stimulus from the preceding CS was lengthened over the range of 2.5 to 10 sec. The results were discussed in terms of the interaction of short-term memory representations with subsequent stimulation, and a specific theory concerning the effects of “priming.”     

A surprising extension of the preconditioned stimulus beyond the cotermination of the US and the added element does not alleviate blocking to the added element

Rats were subjected to the Kamin two-stage blocking procedure. First, a stimulus, A, was conditioned and then reinforced in compound with a target stimulus, B. During compound training, an attempt was made to alleviate blocking of conditioning to B by presenting a “surprise” stimulus for 5 sec following the reinforced AB compound. The surprise stimulus consisted of the continued presentation of the previously reinforced element, A. During A training, A had never extended beyond the moment of reinforcement; thus, its extension beyond reinforcement during AB training was hypothesized to be “surprising” and, therefore, expected to alleviate blocking. Blocking in this condition was, however, just as strong as in a standard blocking condition. The results do not favor a surprise interpretation of unblocking but do seem to be consistent with other theoretical views (e.g., Rescorla & Wagner, 1972).     

Blocking of the sexual conditioning of differentially effective conditioned stimulus objects

The blocking phenomenon was investigated in the sexual response system of male Japanese quail. Access to a live female quail served as the unconditioned stimulus (US). The same audiovisual cue served as the pretrained stimulus in all of the experiments. Following asymptotic conditioning of the audiovisual cue, a second conditioned stimulus (CS2) was added. In Experiment 1, CS2 was a rectangular wood block that had little or no resemblance to a female quail and could not support copulatory behavior. In Experiment 2, CS2 was a terrycloth object that had no quail parts but could support copulatory behavior, and, in Experiment 3, CS2 was a terrycloth object that had a taxidermically prepared head of a female quail added. The terrycloth-only object supported more rapid conditioning than did the wood block, but the blocking effect was obtained with both kinds of stimuli. Approach responding to the terrycloth + head object required pairing it with copulatory opportunity, and the terrycloth + head object supported at least as rapid conditioning as did the terrycloth-only object. However, responding to the terrycloth + head object was not blocked by the pretrained audiovisual cue. These results indicate that the blocking effect occurs in sexual conditioning even with stimulus objects that can support copulation. However, the addition of species-typical head cues to an object makes that object such a powerful stimulus that conditioned approach responding to it cannot be blocked by a previously conditioned arbitrary audiovisual cue.     

One-trial simultaneous and backward excitatory fear conditioning in rats: Lick suppression, freezing, and rearing to CS compounds and their elements

Three experiments with rat subjects sought to enhance one-trial excitatory simultaneous and backward fear conditioning by using a two-element compound conditioned stimulus (CS) instead of only a single element. During conditioning, experimental groups received a 4-sec CS either coextensively with a 1-mA grid-shock unconditioned stimulus (US) or immediately after US termination. In subsequent tests, CSs evoked more lick suppression and freezing in these groups than in various controls. Compound CSs evoked more lick suppression and freezing than did CS elements, but did so equally for experimental and control groups. Therefore, the use of compounds did not enhance conditioning. Unexpectedly, an explicitly unpaired control in which CS followed US termination by 3 min tended to show more CS-evoked suppression and freezing than did a control in which CS preceded US onset by 3 min. This result raises the possibility that associations between the CS and the training context might engender responding to backward-paired CSs.     

Classical conditioning in paramecia

SingleParamecium caudatum were conditioned by pairing ac-generated electric shock (US) with a vibratory stimulus (CS) produced by an auditory speaker. Naive paramecia subjected to shock reliably exhibited a backwards jerk and axial spinning similar to the avoiding reaction described by Jennings in 1904. Such responses did not occur initially to CS alone, but increasingly appeared during the CS period preceding shock pairing (delayed conditioning paradigm). Control subjects given the CS and UCS at the same intervals, but explicitly unpaired, did not show a sustained increase of responses to the CS alone. Short-term memory was demonstrated by subjects first conditioned and then presented CS alone during extinction. These subjects were readily reconditioned. Paramecia trained and stored for 24 h showed reliable memory savings as compared to stored control subjects. Other paramecia were differentially conditioned by training with two CSs. Following the recommendations of Rescorla (1967), a procedure was designed for truly random presentation of the CS and UCS as an additional control for pseudoconditioning. Single paramecia were conditioned with intervals between CSs randomly ranging from 8 to 32 sec. Control subjects received the same number of CSs and UCSs, which were administered independently and randomly during the same total session duration. Thus, CS and UCS were occasionally paired for control subjects. The responses to CS in the conditioned group were anticipatory conditional responses due to the pairing contingency and not wholly due to pseudoconditioning.     

Signal-directed behavior in the rat: Interactions between the nature of the CS and the nature of the UCS

Two experiments are described, which involved the investigation of interactions between the nature of the conditioned stimulus (CS) and the nature of the unconditioned stimulus (UCS) in producing signal-centered behavior. In Experiment 1, rats received response-independent heat reinforcement in a cold environment. For some groups, this heat UCS was signaled by presentations of a standard aluminum retractable lever; for other groups, it was signaled by a retractable lever covered in acrylic fur (furry lever CS). Only the subjects that received the furry lever CS paired with heat exhibited differential CS-contact behavior, when compared with unpaired, aluminum lever, and warm control subjects. In Experiment 2, hungry rats received pairings of either an aluminum or a furry lever with food (UCS). When compared with unpaired controls, only the subjects that received the aluminum lever paired with food showed differential signal-directed behavior; the subjects receiving the furry lever CS did not show differential contact with the CS, but instead exhibited differential food tray entry behavior during CS presentation. In the two studies, the signal-directed behavior exhibited by subjects resembled either thermoregulatory or feeding behaviors characteristic of rats. The results suggest that signal-directed behavior is determined by a complex interaction between the ecological relevance of the CS and the nature of the UCS—an interaction that can best be described in terms of a behavior systems model of conditioned responding.     

One-trial simultaneous and backward fear conditioning as reflected in conditioned suppression of licking in rats

In three experiments, groups of albino rats received one strictly simultaneous pairing of a 4-sec auditory conditioned stimulus (CS) and a 4-sec 1-mA shock unconditioned stimulus (US). Other groups received a backward pairing, in which the US began before the CS, or a forward pairing, in which the CS began before the US. Control groups received only the US or received both the CS and the US but widely separated in time. Later, the CS was presented while the rats licked a drinking tube for water, and CS-elicited suppression of licking was taken as an index of the Pavlovian conditioned response (CR). It was found that groups receiving a single forward or a single simultaneous pairing suppressed more than groups that had received a backward pairing; and the backward groups, in turn, suppressed more than the control groups. It appears, then, that excitatory fear conditioning, as reflected in conditioned suppression of licking in rats, can be produced in a single trial by both backward and simultaneous conditioning procedures.     

The role of signaled periods of nonreinforcement in responding on a random schedule in autoshaping

Little responding develops to a conditioned stimulus (CS) that is placed in a random relation to an unconditioned stimulus (US). However, if the USs not preceded by that CS are themselves signaled by another stimulus, then the CS does come to elicit responding. This result has been attributed (e.g., by Durlach, 1983) to the signal’s blocking of conditioning to background cues that otherwise would prevent conditioning of the CS. However, Goddard and Jenkins (1987) have suggested the alternative that signaling the USs promotes responding due to the adventitious creation of periods of signaled nonreinforcement. Two experiments were conducted to assess this alternative, involving an autoshaping preparation in pigeons. In Experiment 1, little responding to a keylight CS presented in a random relation to a food US occurred, despite the explicit presentation of a discrete noise signaling periods of no food in the intertrial interval (ITI). Experiment 2 was designed to replicate the procedure of Goddard and Jenkins, in which an auditory stimulus extended throughout the ITI of a random schedule, terminating only prior to extra USs and during the CS. Contrary to their findings, little responding developed to the target CS. However, responding did develop when the sound-free period occurred only prior to the extra USs. These results offer little support for the hypothesis that signaled periods of nonreinforcement promote responding on random schedules. However, they are consistent with the view that signaling of ITI USs acts by preventing conditioning of potentially competitive background cues.     

Context effects on conditioning,extinction, and reinstatement in an appetitive conditioning preparation

Three experiments with rat subjects examined the effects of contextual stimuli on performance in appetitive conditioning. A 10-sec tone conditioned stimulus (CS) was paired with a food-pellet unconditioned stimulus (US); conditioning was indexed by the observation of headjerking, a response of the rat to auditory stimuli associated with food. In Experiment 1, a context switch following initial conditioning did not affect conditioned responding to the tone; however, when the response was extinguished in the different context, a return to the original conditioning context “renewed” extinguished responding. These results were replicated in Experiments 2 and 3 after equating exposure to the two contexts (Experiment 2) and massing the conditioning and extinction trials (Experiment 3). The results of Experiment 1 also demonstrated that separate exposure to the US following extinction reinstates extinguished responding to the tone; this effect was further shown to depend at least partly on presenting the US in the context in which testing is to occur (Experiments 2 and 3). Overall, the results are consistent with previous data from aversive conditioning procedures. In either appetitive or aversive conditioning, the context may be especially important in affecting performance after extinction.     

Context sensitivity of conditioned suppression following preexposure to the conditioned stimulus

Conditioned suppression in rats is often unaffected when the context (or set of background stimuli) is changed following conditioning. This suggests that responding to the conditioned stimulus (CS) can be relatively insensitive to the context in which the CS is presented. In two experiments, we examined whether sensitivity to contextual stimuli is affected by preexposure to the CS. In Experiment 1, when the CS was novel at the outset of conditioning, conditioned suppression was not affected when the context was changed following conditioning. However, when the CS had been preexposed, responding was weaker when extinction occurred outside of the conditioning context. In Experiment 2, responding was again sensitive to the test context, regardless of whether preexposure occurred in the conditioning context or in an alternate context. These results suggest that the extent to which responding is sensitive to context can depend on the conditioning history of the CS.     

Applying Konorski’s model of classical conditioning to signal-centered behavior in the rat: Some functional similarities between hunger CRs and sign-tracking

Two experiments were conducted to investigate functional similarities between “hunger CRs” of Konorski’s (1967) model of appetitive classical conditioning and sign-tracking behavior in rats. Konorski’s model predicts that hunger CRs will be facilitated (1) when a nonrein-forced stimulus similar to the reinforced CS is introduced, and (2) when some CS presentations are unexpectedly nonreinforced. In Experiment 1, hungry rats acquired a leverpress response to a retractable lever that was paired with response-independent food. Following this training, a second lever was introduced whose presentation was not followed by food. The effect of the presence of this second lever was to facilitate responding to the original lever. In Experiment 2, single-lever autoshaping training was followed by a shift from 100% pairing of the lever with food to only 50% of the lever presentations being followed by food. The introduction of partial reinforcement produced an immediate and durable increase in leverpressing. The findings of both experiments are consistent with predictions from Konorski’s model of classical conditioning if sign-tracking is considered as a “hunger CR.”     

Divergence of conditioned eyeblink and conditioned fear in backward Pavlovian training

Two experiments of Pavlovian conditioning with rabbits evaluated the effects of initiating or continuing a conditioned stimulus (CS) after a paraorbital unconditioned stimulus (US). In Experiment 1, backward pairings, in which a CS came on after the US, produced a CS that appeared inhibitory on a measure of eyeblink conditioning but excitatory on a potentiated-startle measure of conditioned fear. In Experiment 2, extending the duration of a CS that came on prior to the US, so that it continued after the US, decreased eyeblink conditioned responses, whereas it increased conditioned fear. The data from the two experiments confirm and extend those of Tait and Saladin (1986), supporting the suppositions of AESOP (Wagner & Brandon, 1989) that conditioned eyeblink and conditioned fear can be dissociated under various temporal relationships between the CS and US.     

The influence of the information value provided by prior-cuing treatment on the reactivation of memory in preweanling rats

In two experiments, the influence of exposure to a CS? on the acquisition and retention of a conditioned odor aversion was examined. Preweanling rats were given exposure to the CS? either prior to (CS?/CS+) or following (CS + /CS?) the pairing of a second odor (the CS+) with footshock. The results of Experiment 1 indicated that subjects in both of the treatment conditions acquired aversions of comparable strength to the odor paired with footshock and that retention of the odor aversion was not affected by order of stimulus presentation during conditioning. Experiment 2 indicated, however, that the effectiveness of pretest exposure to various elements of the conditioning episode in reactivation of the memory for conditioningwas dependent on the order of stimulus presentation during conditioning. This differential effectiveness of the various reactivation treatments is discussed in terms of their relationship to the associative “status” of the stimuli present during conditioning and in terms of the information provided to the animal by the reactivation treatment.     

Secondary extinction in Pavlovian fear conditioning

Pavlov (1927/1960) reported that following the conditioning of several stimuli, extinction of one conditioned stimulus (CS) attenuated responding to others that had not undergone direct extinction. However, this secondary extinction effect has not been widely replicated in the contemporary literature. In three conditioned suppression experiments with rats, we further explored the phenomenon. In Experiment 1, we asked whether secondary extinction is more likely to occur with target CSs that have themselves undergone some prior extinction. A robust secondary extinction effect was obtained with a nonextinguished target CS. Experiment 2 showed that extinction of one CS was sufficient to reduce renewal of a second CS when it was tested in a neutral (nonextinction) context. In Experiment 3, secondary extinction was observed in groups that initially received intermixed conditioning trials with the target and nontarget CSs, but not in groups that received conditioning of the two CSs in separate sessions. The results are consistent with the hypothesis that CSs must be associated with a common temporal context during conditioning for secondary extinction to occur.     

Massive preexposure and preexposure in multiple contexts attenuate the context specificity of latent inhibition

Latent inhibition, which refers to attenuated responding to a conditioned stimulus (CS) after CS-unconditioned stimulus (CS-US) pairings as a result of CS-alone presentations prior to the pairings, is often attenuated if preexposure and conditioning occur in different contexts (i.e., it is context specific). Here we report two conditioned lick suppression experiments, using rat subjects, that examined whether manipulations known to attenuate the context specificity of extinction could also eliminate the context specificity of latent inhibition. Context specificity of latent inhibition was eliminated when the CS was preexposed in multiple contexts (Experiment 1) and when the CS was massively pre-exposed in the training context alone (Experiment 2). These results and their practical implications are discussed in the framework of contemporary theories of latent inhibition.     

Dissociation of conditioned appetitive and consummatory sexual behavior: Satiation and extinction tests

Sexual responses were conditioned in male Japanese quail using the opportunity to copulate with a female as the unconditioned stimulus (US). The conditioned stimulus (CS) was a 3-D object made of a taxidermically prepared female quail head mounted on a terry-cloth body. Both appetitive conditioned responses (approach and proximity to the CS) and consummatory conditioned responses (mount and cloacal contact directed toward the CS) developed when 2-min presentations of the CS were followed immediately by the US, but not when the CS and US were separated by trace intervals of 10 or 20 min (Experiment 1). Postconditioning sexual satiation suppressed conditioned cloacal contact responses more than conditioned approach to the CS (Experiment 2), and “acute” extinction suppressed both conditioned mounting and conditioned cloacal contact responses more than conditioned approach to the CS (Experiment 3). These results demonstrate a functional dissociation between conditioned appetitive and consummatory responses and imply that the motivational and/or associative mechanisms underlying the two types of behavior are distinct.