Pavlovian counterconditioning: changing the suppressive properties of shock by association with food.

Two experiments are described in which rats received a series of shock and food presentations in a Pavlovian counterconditioning sequence. Subsequently, the capacity of the shock to act as a reinforcer during conditioned emotional response (CER) training was assessed. In the first experiment, following each shock during counterconditioning by a food presentation was found to retard the development of suppression during CER training relative to control conditions in which different groups of rats received either a sequence of explicitly unpaired shock and food presentations, shock presentations alone, or food presentations alone. The second experiment demonstrated that when the magnitude of the food presentations is held constant, the attenuation of the suppressive properties of shock by pairing with food depends on the magnitude of the shock. The results are interpreted as demonstrations of Pavlovian counterconditioning, which, it is suggested, modifies the general aversiveness of a noxious stimulus.     

The effects of lateral, medial, or complete septal lesions on positive conditioned suppression in rats

This study investigated the effects of lateral, medial, or complete septal lesions in rats on lever pressing and US-approach behaviors during the presentation of a light followed by two free food pellets. Ten days after surgery, groups of rats received 20 sessions of a random interval (RI) 60-s schedule of food reinforcement. The positive conditioned suppression paradigm consisted of 20 sessions of a 10-s light paired with free food while the rats were responding on the RI 60-s schedule. All groups of rats significantly suppressed lever pressing during the 10-s light presentation. During the last four sessions of the 10-s light condition, rats with lateral septal lesions had a significant increase in food-tray entries during the light presentation, while the other groups decreased lever pressing without a change in food-tray entries during the 10-s light presentation. Although rats with septal damage generally have difficulty inhibiting responses in a variety of operant situations, rats with lateral, medial, or complete septal lesions showed no impairment in positive conditioned suppression. This study also suggests that when the spatial arrangement of the CS and the type of US were manipulated to minimize sign-tracking and goal-tracking behaviors, emotional reactions during the presentation of the CS must be considered a factor influencing positive conditioned suppression.     

Counterconditioning of appetitive and defensive CRs in rabbits

Two experiments examined interactions between conditioned appetitive and defensive responses in the rabbit. In Experiment I, a conditioned jaw-movement response was established by following presentations of a clicker CS with intra-oral sucrose delivery on 50% of trials. The jaw-movement response was then maintained on this partial reinforcement schedule during a counterconditioning phase. A group which received para-orbital shock paired with the CS on non-sucrose trials showed acquisition of eyeblink responding and suppression of jaw-movement responding to the CS, in comparison to control groups which received either no shock or unpaired presentations of the CS and shock. Experiment II was identical in design to Experiment I except that the roles of the sucrose and shock reinforcers were reversed. The paired group acquired a conditioned jaw-movement response when sucrose was added in the counterconditioning phase, but in contrast to Experiment I showed a slight enhancement of the previously established eyeblink response. The asymmetry of appetitive and defensive counterconditioning was discussed in relation to opponent-process theories of motivation and reinforcement.     

Preference and response suppression under different correlations between shock and a positive reinforcer in rats

A lever-press response by rats was reinforced by food in two successively presented types of trial signalled by different discriminative stimuli. When responding was punished by a shock in one type of trial, a groups for which the shock always preceded the reinforcer by .5 sec (positive correlation) showed less suppression in those trials than a group which received the shock and food separated in time (negative correlation). When, in a second experiment, rats were given a choice between food alone in one end of a shuttle box and either positively or negatively correlated shock and food in the other on a concurrent schedule, the group receiving the negatively correlated shock showed a greater preference for the food alone end. On the basis of a third experiment in which a tone was substituted for the shock in the choice situation, it was argued that the effect of correlation was not simply due to the stimulus properties of the shock. A final experiment demonstrated that when shock punishment is administered during extinction of the lever-press response, the rate of extinction is slower if the shock has been previously paired with the food reinforcer. Pairing a shock with food seems to attenuate the intrinsic aversiveness of the shock through Pavlovian counterconditioning.     

Appetitive-aversive interactions: Superconditioning of fear by an appetitive cs

Four groups of rats received conditioned suppression training in which a tone and light compound was reinforced with shock. If the light had been previously paired with free food, enhanced fear conditioning accrued to the tone during compound training relative to control groups pre-exposed to the light alone, the light semi-randomly associated with food, or the light unpaired with food. The second experiment replicated the difference in aversive conditioning for the groups receiving the light either paired or unpaired with food. The results are discussed in terms of the functional similarity of a conditioned excitor for food and a conditioned inhibitor for shock.     

A modified counterconditioning procedure prevents the renewal of conditioned fear in rats

Two studies using an ABA design examined the Extinction and renewal of conditioned barpress suppression. Following lights-off and foot shock pairings in Context A, rats were placed in Context B and were given either a standard counterconditioning procedure where the lights-off CS was paired with a novel food US delivered freely or a modified counterconditioning procedure where CS-US pairings only occurred if the rat earned the US by performing a required behavior during the CS. Results indicated that the modified counterconditioning procedure thwarted ABA renewal but the conventional counterconditioning procedure did not reduce ABA renewal any more than nonreinforced exposure to the CS alone. Furthermore, the response required during the modified counterconditioning procedure could be one used as a baseline response during Acquisition of fear or it could be a novel response. Implications of the results for theories of Extinction and renewal of fear are discussed.     

One-trial conditioned suppression: effects of instructions on extinction

Two experimental groups of undergraduate volunteers received a single Pavlovian conditioning trial consisting of a paired presentation of a tone conditioned stimulus (CS) and a shock unconditioned stimulus (UCS). Control groups received either the CS alone or the UCS alone. Subjects from one experimental group were subsequently instructed that they would not receive further shocks, while the other experimental group received no such instructions. The CS alone was then presented once to all four groups while subjects were engaged in a button-pressing task maintained by slide reinforcement. During the latter phase, rate of button-pressing was measured. Classically conditioned suppression of button-pressing was obtained in the noninstructed experimental group but not in the instructed group. The results demonstrate that suppression can be a sensitive index of Pavlovian conditioning in humans but question the use of conditioned suppression as an adequate experimental analog of clinically observed anxiety-motivated behavior.     

Autoshaping a leverpress in rats with lateral, medial, or complete septal lesions

Rats with either control operations or lateral, medial, or complete septal lesions received 600 trials of leverpress training using an autoshaping procedure, i.e., food delivery followed a 10 s illuminated lever presentation, or occurred immediately after a leverpress. Rats with complete septal lesions acquired the leverpress faster than controls and had more food-tray entries per minute during the first 100 trials than the other groups. Rats with lateral or medial septal lesions had leverpress and food-tray entries equivalent to controls. The facilitation of autoshaping a leverpress may partially be explained by the general increase in motor reactivity to stimuli found following septal lesions.     

Associations in Pavlovian conditioned inhibition

Two experiments with rat subjects used a variety of transfer tests to examine the associations learned when Pavlovian inhibition is established by an A+, AX− paradigm. Experiment 1 found in a conditioned suppression situation that inhibition conditioned to X with one exciter (A) readily transferred to another exciter (B) which had been paired with the same shock US. Transfer occurred even when the response to A had been extinguished prior to testing with B. However, X did not inhibit a general activity response produced by a B which had been subsequently paired with a food US. Experiment 2 employed a Pavlovian conditioning situation in which A and B, when separately paired with the same food US, evoked dissimilar responses. Nevertheless, an inhibitor trained in an A+, AX− paradigm successfully inhibited the different response evoked by B. However, such an X did not inhibit the behaviors acquired by A or B when they were subsequently paired with a shock US. The transfer of Pavlovian inhibition across conditioned stimuli and responses but not across unconditioned stimuli is consistent with the notion that a conditioned inhibitor acts to prevent activation of a US representation which would normally be activated by conditioned exciters.     

Time-dependent changes in conditioned suppression

Time-dependent changes in a response following aversive conditioning were investigated using a conditioned suppression procedure in a within-subjects design. Four groups of pigeons received Pavlovian conditioning “off the baseline”, immediately followed by an operant task. During the Pavlovian phase, two groups received a forward pairing of a tone with shock, one group received a backward pairing, and one group received a truly random pairing. One of the forward pairing groups also received a delay between the Pavlovian and operant phases. For all groups, key pecking was reinforced on a variable-interval schedule during the operant phase. Testing sessions were identical to training sessions, except that the tone used during Pavlovian conditioning was presented either 0, 15, 30, 45, of 60 minutes after the operant phase began. Testing sessions in which the Pavlovian phase was omitted were also included. The results showed suppression to change as a function of the retention interval, with maximum suppression occurring at intermediate intervals. This U-shaped function was obtained for 11 of the 12 pigeons in the forward-pairing groups and for three of the five in the truly random group. Pigeons in the background pairing group did not show changes in suppression as a function of the retention interval.     

Conditioned suppression of an avoidance response by a stimulus paired with food

Three food-deprived Long-Evans rats were exposed to a non-discriminated shock avoidance procedure. Superimposed upon this operant avoidance baseline were periodic presentations of a conditioned stimulus that was paired with food, the unconditioned stimulus. These pairings resulted in increases in the rate of shock over that recorded when the conditioned stimulus was not present. A traditional suppression ratio failed to reveal any differential effect of the conditioned stimulus on the overall rate of avoidance responding, although all subjects showed a consistent pattern of pausing and postshock response bursts during presentations of the conditioned stimulus. When food was withheld during a final extinction phase, the conditioned stimulus ceased to occasion increases in shock rates and disruptive postshock response bursts were eliminated. An analysis of conditioned suppression procedures is proposed that stresses not only operant-Pavlovian or appetitive-aversive incompatibility, but also the manner in which the baseline schedule of reinforcement affects operant behavior changes that are elicited by the superimposed Pavlovian procedure.     

Hippocampal associative cellular responses: dissociation with behavioral responses revealed by a transfer-of-control technique

Multiunit activity was recorded in the CA3 field of the dorsal hippocampus in freely moving rats during classical conditioning and subsequent presentation of the CS on operant baselines for food reward as well as shock avoidance. Rats were first trained in a nonsignaled bar-pressing-dependent shock omission task and in a food-motivated lever-pressing task (60-s VI). Five sessions with presentations of a previously habituated tone as a CS paired with footshock as a US were then given. Testing was carried out by presenting the CS alone while behavioral responses were maintained by reinforcement in both instrumental tasks on alternate sessions. As expected, the CS induced a marked suppression of lever pressing for food reward and a marked enhancement of bar-pressing for shock avoidance. The analysis of the frequency of multiunit discharges to the CS revealed that the hippocampal cellular responses established during classical conditioning were maintained while two different behavioral responses were exhibited to the CS. The results showed that the associative response of hippocampal neurons may be dissociated from the Pavlovian conditioned responses the CS elicits. They support the hypothesis that hippocampal cellular responses represent a neural index of the acquired CS-US associative representation.     

Constraints on Pavlovian conditioning of the pigeon: Relative conditioned reinforcing effects of red-light and tone CSs paired with food

Recent Pavlovian conditioning experiments presented all possible CS-US combinations of red-light and tone CSs and food and shock USs to separate groups of pigeons. Pigeons receiving shock USs demonstrated conditioned head raising followed by prancing to both CSs, but CRs were acquired more rapidly to tone than to red light. Although pigeons receiving food USs rapidly acquired a conditioned response of pecking to the red-light CS, there was no evidence of conditioned responding in groups receiving tone-food pairings. This outcome left open the possibility that Pavlovian pairings of tone and food may have resulted in association formation that was not revealed in performance. The present series of experiments attempted to reveal that association, using an indirect method of assessment, conditioned reinforcement. Experiment 1 demonstrated that both red light and tone paired with food became positive conditioned reinforcers, suggesting that an association between tone and food was formed in the same number of Pavlovian conditioning trials that previously failed to yield any direct evidence of conditioning. Experiment 2, which presented fewer conditioning trials, revealed that the tone-food association was formed less rapidly than the red light-food association. Experiment 3 demonstrated that the observed outcomes were not attributable to unconditioned, rather than conditioned, reinforcing effects of the Css.     

Learned fear of "unseen" faces after Pavlovian, observational, and instructed fear

This study compared fear learning acquired through direct experience (Pavlovian conditioning) and fear learning acquired without direct experience via either observation or verbal instruction. We examined whether these three types of learning yielded differential responses to conditioned stimuli (CS+) that were presented unmasked (available to explicit awareness) or masked (not available to explicit awareness). In the Pavlovian group, the CS+ was paired with a mild shock, whereas the observational-learning group learned through observing the emotional expression of a confederate receiving shocks paired with the CS+. The instructed-learning group was told that the CS+ predicted a shock. The three groups demonstrated similar levels of learning as measured by the skin conductance response to unmasked stimuli. As in previous studies, participants also displayed a significant learning response to masked stimuli following Pavlovian conditioning. However, whereas the observational-learning group also showed this effect, the instructed-learning group did not.     

Counterconditioning and exposure-only in the treatment of specific (conditioned suppression) and generalized fear in rats

In Experiment 1 the conditioned suppression technique was used to condition specific fear, suppression of operent lever pressing for food to a discrete CS. The efficacy of four treatment conditions on fear reduction was evaluated. Counterconditioning in which exposure to the CS was contiguously paired with food was significantly less effective than noncontiguous CS exposure and food. An exposure-only effect was indicated by the superiority of all three treatments involving CS exposure (the above two plus a typical conditioned suppression extinction procedure) to treatment consisting of food only. The reverse counterconditioning effect and the exposure effect are consistent with current views that emphasize the centrality of aversive stimulus exposure in fear reduction. Experiment 2 investigated elimination of generalized fear produced by unsignalled, inescapable shocks in the lever-pressing apparatus. Two treatments (counterconditioning and exposure-only) were equally effective and they were superior to no exposure control treatment. The results of the two experiments reinforce recent attempts toward a reevaluation of the role of anxiety-competing responses in elimination of fear.     

Lesions of Periaqueductal Gray Dissociate-Conditioned Freezing From Conditioned Suppression Behavior in Rats

It is commonly assumed that suppression of an ongoing behavior is an indirect measure of freezing behavior. We tested whether conditioned suppression and freezing are the same or distinct conditioned responses. Rats were trained to press a bar for food and then given fear-conditioning sessions in which a tone was paired with a foot shock (two pairings a day for 2 days). They then received either sham or electrolytic lesions of the periaqueductal gray (PAG). Post-training PAG lesions blocked freezing to the conditioned stimulus (CS), but had no effect on the suppression of operant behavior to the same CS. Thus, conditioned suppression and freezing, which both cause a cessation in activity, appear to be mediated by separate processes.     

Positive conditioned suppression: conditioned suppression using positive reinforcers as the unconditioned stimuli

Research has revealed the phenomenon of conditioned suppression in which the rate of responding is reduced during a stimulus that is paired with noncontingent shock. The present study replicated this procedure, but used noncontingent positive reinforcers instead of the aversive shock. The lever-pressing responses of rats were reinforced with food or water. While the rats were responding, a stimulus was occasionally presented and paired with the delivery of a noncontingent positive reinforcer, which was either food, water, or brain stimulation for different rats. The result was a reduction in the rate of responding during the conditioned stimulus. This finding shows that conditioned suppression occurs during a signal for reinforcing as well as aversive stimuli.     

Acquired equivalence and distinctiveness of cues

In Experiments 1 and 2, rats received initial training in which two stimuli (A and N) were either followed by the same consequence (food) or by different consequences (food and no food). Subsequently N was paired with electric shock and the generalization of conditioned suppression to A was assessed. Suppression to A was more marked when A and N had both been followed by food than when they had had different outcomes. In Experiment 3, 3 stimuli (A, B, and N) were presented initially. For one group, A and N were paired with food and B was nonreinforced: for a second group, B was paired with food and A and N were nonreinforced. Generalization of suppression was found to be more substantial to A than to B for both groups. These results indicate that the extent to which stimuli are treated as being equivalent is partly determined by their reinforcement histories.     

Signal duration and suppression of operant responding by free reinforcement

Pigeons were trained on a VI (variable interval) schedule of food presentation with a superimposed schedule of response-independent food. Substantial suppression of the operant response rate occurred when the free food was presented without a signal. When the free food was preceded by a short (4 sec) signal, the degree of suppression was similar to that with unsignaled free food. But when the signal was lengthened to 12 sec, the degree of suppression was substantially reduced. Experiment 3 assessed the effect of signal duration using a baseline schedule of delayed reinforcement, in which contingent reinforcers were themselves preceded by a signal. The signal preceding the free reinforcers was then either the same as or different from this contingent signal. Signal duration effects occurred only when the two types of signals were different. These differences as a function of signal duration have implications for both “context-blocking” and “comparator” interpretations of the effects of noncontingent reinforcement in both Pavlovian and operant procedures.     

Shock as a signal for shock or no-shock: a feature-negative effect in conditioned suppression.

Rats were trained in conditioned suppression discriminations where shock at the beginning of a trial signaled either shock or no-shock at the end of the trial. In the shock-positive condition, shock at the beginning of a presentation of white noise signaled that noise would end with shock; noise that did not begin with shock did not end with shock. In the shock-negative discrimination, shock at the beginning of noise signaled that noise would not end with shock; presentations of noise that did not begin with shock ended with shock. In shock-random training, shock at the beginning of noise did not reliably signal whether the noise presentation would or would not end with shock. Most subjects in shock-negative training quickly developed a differential pattern of suppression on positive (shock reinforced) trials and no suppression on negative (nonreinforced) trials. The shock-positive discrimination was much more difficult to establish and was not acquired by the majority of the rats. This "feature-negative" effect is a clear exception to the general superiority of feature-positive learning commonly observed in discriminations based on a single distinguishing feature. The results are discussed in terms of Pavlovian stimulus-shock contingencies in the shock-positive and shock-negative paradigms, which appear to favor rapid development of the shock-negative discrimination.