Retardation of conditioned excitation following operational inhibitory blocking

Water-deprived adult rats were used in a conditioned-suppression-of-licking procedure to determine the effect of inhibitory training with a novel stimulus trained in simultaneous compound with a previously established conditioned inhibitor. This procedure constitutes an inhibitory analogue to the excitatory blocking procedure in classical conditioning. The conditioned-inhibition training consisted of either explicitly unpaired CS and US presentations or negative contingency training, in which the likelihood of the US was greater in the absence than in the presence of the CS, but the CS and the US were occasionally paired. To assess conditioned inhibition, a retardation test was used, and comparable retardation was obtained for subjects that were administered the blocking treatment and control subjects given similar conditioned-inhibition training with a compound stimulus in which the nontarget element was not previously established as a conditioned inhibitor.     

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.     

Enhancement of conditioned inhibition via an extinction treatment

Rats were used in a conditioned-suppression paradigm to determine whether an extinction treatment would enhance a moderately developed conditioned inhibitor (CS?). To dissipate unconditioned suppression to the training stimuli, the subjects were first habituated to the stimuli and then given Pavlovian conditioned-inhibition (CI) training involving reinforced presentations of a clicker and nonreinforced compound presentations of that stimulus and the intended CS?, either a light or a tone. Thereafter, experimental subjects received presentations of their CS? by itself, whereas controls received no further training. Following the occurrence and loss of conditioned suppression to the CS? in the extinction phase, summation and retardation tests showed enhanced CI for the experimental subjects relative to both the controls and their own earlier levels of inhibitory performance. In fact, the enhanced inhibition for the experimental subjects approximated that shown by a comparison group for which the CS? had been strongly developed as an inhibitor. These findings suggest that an excitatory representation is associated with the CS? early in CI training, and that subsequent presentations of the CS? by itself strengthen its inhibitory effect by allowing it to be nonreinforced in the presence of that representation.     

Renewal of Pavlovian conditioned inhibition

Prior research has demonstrated renewal, which is the ability of contextual cues to modulate excitatory responding to a Pavlovian conditioned stimulus (CS). In the present research, conditioned lick suppression in rats was used to examine similar contextual modulation of Pavlovian conditioned inhibition. After Pavlovian conditioned inhibition training with a CS in one context, subjects were exposed to pairings of the CS with an unconditioned stimulus (US) either in the same or in a second context. Results indicated that, when the CS was paired with the US in the second context, the CS retained its inhibitory control over behavior, provided that testing occurred in the context used for inhibition training. However, when the CS-US pairings occurred in the inhibition training context, the CS subsequently proved to be excitatory regardless of where testing occurred. These observations indicate that conditioned inhibition is subject to renewal.     

Summation of responding to CSs and an excitatory test context

The influence of test-trial delay of CS onset in obtaining response summation of an excitatory CS and an independently conditioned context was investigated. Water-deprived rats were given tone-shock and click-shock pairings in the training context and unsignaled footshocks in the test context. Durations of lick suppression in response to the tone (Test 1) and the click (Test 2) were then assessed in the test context. Licking was more suppressed when CS onset occurred early in the test session (e.g., 0 sec) than when it occurred later in the test session (e.g., 300 sec). The results from control groups that had received shock in an irrelevant context rather than the test context indicated that this effect was due to fear of the test context rather than diffuse, nonassociative fear. With onset of the clicks early in the test session on the second test day, response summation of the test context with the clicks was observed. This suggests that appreciable spontaneous recovery had occurred from any test-context extinction that took place on the first test day. We conclude that early onset of the CS on the test trial favors response summation of CSs and test context.     

Preweanling and adult rats treat conditioned light-tone combinations differently

Ontogenetic differences in processing light-tone compounds were discovered in preweanling (17-day-old) and adult (60–80-day-old) rats. Suppression of general activity was used as an index of the magnitude of conditioned fear following a single training session in which a CS+ was paired with mild footshock. In Experiment 1, rats were trained on discriminations in which the CS? consisted of a light and the CS+ was either a tone alone (simple discrimination) or a light-tone compound (simultaneous feature-positive discrimination). Adults and preweanlings given each type of discrimination were then tested for fear of the CS? and a target stimulus (tone alone or light-tone compound). Adults in all groups displayed greater fear of the target than of the CS?. Preweanlings, however, discriminated the CS? from the target only when the target was the same as the original CS+. Experiment 2 revealed that age-related differences in conventional stimulus generalization is not a likely explanation for the pattern of results found in Experiment 1. Experiment 3 revealed age-related differences in expressed fear of a serial feature-positive discrimination; adults, but not preweanlings, showed greater fear of the compound than of the CS?. Alternative interpretations of the results from these experiments are discussed, and the general conclusion is that adults appear more inclined to process elements of a compound stimulus selectively, whereas preweanlings seem more likely to process the compound unselectively, with roughly equivalent processing of each element.     

Temporal coding in conditioned inhibition: Retardation tests

Two lick suppression experiments with rats were conducted in order to determine the nature of the temporal information that is encoded as a result of Pavlovian conditioned inhibition training (conditioned stimulus {CS} A→unconditioned stimulus {US}/AX→noUS). After inhibition training, the conditioned inhibitor (X) was paired with the US in order to measure inhibition, as assessed through retarded behavioral control by CS X. Three temporal relationships were manipulated: the A-US interval, the X-A interval of inhibition training, and the X-US interval of the retardation test pairings. Retardation was greatest when the X-US temporal relationship matched the time at which the US was expected (but not delivered) on the X-A inhibition training trials. Thus, the present experiments provide evidence with retardation tests that, during conditioned inhibition training, subjects encode the temporal location of the omitted US relative to the inhibitory CS. These findings complement those of previous studies using summation tests of conditioned inhibition (Barnet & Miller, 1996; Denniston, Blaisdell, á Miller, 1998; Denniston, Cole, & Miller, 1998).     

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.     

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.     

Overshadowing and CS duration: counteraction and a reexamination of the role of within-compound associations in cue competition

In the present experiments, we examined the role of within-compound associations in the interaction of the overshadowing procedure with conditioned stimulus (CS) duration, using a conditioned suppression procedure with rats. In Experiment 1, we found that, with elemental reinforced training, conditioned suppression to the target stimulus decreased as CS duration increased (i.e., the CS duration effect), whereas, with compound reinforced training (i.e., the overshadowing procedure), conditioned suppression to the target stimulus increased as CS duration increased. In subsequent experiments, we replicated these findings with sensory preconditioning and demonstrated that extinction of the overshadowing stimulus results in retrospective revaluation with short CSs and in mediated extinction with long CSs. These results highlight the role of the duration of the stimulus in behavioral control. Moreover, these results illuminate one cause (the CS duration) of whether retrospective revaluation or mediated extinction will be observed.     

Conditioned inhibition training of the competing cue after compound conditioning does not reduce cue competition

In each of two experiments, we studied Pavlovian fear conditioning (as assessed by barpress conditioned suppression) in 32 albino rats. Following a two-stage cue-competition procedure (A+ then AX+), we subjected the competing cue (A) to conditioned inhibition training (B+, BA?) before testing the target cue (X). Conditioned inhibition training was designed to weaken the putative A-unconditioned stimulus (US) association, perhaps changing it to an A-no-US association. Performance-deficit theories of cue competition, such as comparator theory and retrieval-interference theory, predict that such procedures should weaken cue competition, causing Conditioned Stimulus X (CS X) to evoke strong responding. The same prediction can be deduced from recent acquisition-focused models (Dickinson &; Burke, 1996; Van Hamme &; Wasserman, 1994). In opposition to this prediction, however, we found in both experiments that conditioned inhibition training had no detectable effect on cue competition even though it successfully abolished conditioned responding to CS A. In Experiment 2, moreover, we found evidence against the hypothesis that the weak response to CS X was due to generalization decrement rather than to cue competition. Results favor early learning-deficit theories of cue competition over performance-deficit theories and over the recent acquisition-focused models.     

Release from latent inhibition with delayed testing

A conditioned emotional response procedure was used to study the interactive effects of stimulus preexposure and retention interval in rats. In Experiment 1, the subjects were conditioned by presenting a light CS paired with mild footshock as the US. Half of the subjects were given nonreinforced preexposure to the CS, and the others were not. Separate preexposed and nonpreexposed groups were then tested 1,7, or 21 days after conditioning. Suppression of ongoing activity was used to assess the degree of conditioned fear. Latent inhibition was found at the 1-day retention interval; the preexposed subjects displayed less conditioned fear than did the nonpreexposed subjects. In contrast, equally strong conditioned fear was expressed by the preexposed and the nonpreexposed groups tested after the 7- and the 21-day retention intervals. These results indicate a release from latent inhibition similar to that obtained with conditioned taste aversions (Kraemer & Roberts, 1984). The results of Experiment 2 suggest that retention-interval-induced increases in sensitization, pseudoconditioning, or neophobia cannot account for the release from latent inhibition effect obtained in Experiment 1. The implications of these findings for a retrievaloriented view of latent inhibition are discussed.     

Increase in retention of a taste aversion by weanling rats after a long interval

Weanling rats were tested for retention of an aversion to a novel flavor (chocolate milk) that had been conditioned as a single-element conditioned stimulus (CS) or in compound with a novel ambient odor (banana). The presence of the ambient odor during conditioning had no effect on flavor aversion shortly thereafter, confirming previous results. The flavor aversion observed 21 days after conditioning, however, was significantly stronger for pups conditioned with the single-element CS than for those given the flavor-odor compound as the CS. This retention effect was due to a surprisingincrease in the conditioned aversion observed 21 days after conditioning with the single-element CS. A second experiment confirmed this paradoxical increase in retention of the aversion to chocolate milk. This experiment also verified that no such increase occurred in retention of the conditioned aversion to a different flavor (saccharin), whether the initial aversion was strong or weak. The results may be explained in terms of generalized latent inhibition from consumption of mother’s milk.     

Signaling and affective functions in Pavlovian conditioning

The present study employed a Pavlovian-instrumental-transfer paradigm to investigate the role of conditioned fear in appetitive discrimination learning. Each of three Pavlovian training procedures was used to establish a conditioned fear excitor (CS+), a “neutral” CS (CSo), and a conditioned fear inhibitor (CS?). Then, the CSs were administered to rats in the three groups contingent upon the rewarded response in a difficult visual discrimination. In addition, half of each group received shock punishment for each incorrect response. Relative to CSo, CS+ facilitated performance in contrast to the usual interfering effect of conditioned suppressors; conversely, CS? retarded performance even when its reinforcing action (fear inhibition) was potentiated by punishment for the incorrect response. These results, together with other findings showing a reversed outcome when the CSs are administered for the incorrect response, indicate that Pavlovian conditioning comprises both general signaling and affective functions, the former reflecting a basic “expectancy” or nominal type of cognitive processing in the rat.     

Blocking with serial and simultaneous compounds in a trace conditioning procedure

Blocking of conditioned suppression in rats was studied in three experiments using serial and simultaneous compounds in Pavlovian trace conditioning procedures. Experimental groups were first given trace conditioning trials with a 2-sec stimulus (A) presented at least 60 sec before an electric grid shock US. Next, both experimental and control groups received reinforced trials with a compound stimulus (AB). Both A and B were 2 sec in duration and were presented at least 60 sec before the US. For some groups during AB training, the A stimulus preceded the B stimulus; for others, B preceded A; for still others, A and B occurred simultaneously. Conditioning was subsequently assessed separately to both A and B. The results were as follows: First, varying the interval between the onset of A and the US during A training appeared to produce significantly different levels of conditioning to A but did not detectably affect A’s ability to block conditioning to B. Second, blocking was observed in both simultaneous and serial procedures. Third, in the serial procedure, A blocked conditioning to B whether it preceded B or followed B in the AB compound. Fourth, in tests given after AB conditioning, the experimental and control groups suppressed similarly to A. The relevance of these results to the conditioning model of Rescorla and Wagner (1972) and to Mackintosh’s (1975b) theory of attention are discussed.     

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.     

Blocking and enhancement of fear conditioning by appetitive CSs

Prior research on Pavlovian-to-instrumental transfer has shown that when a CS previously associated with shock (AvCS+) is presented contingent upon a choice response to a discriminative stimulus for food reinforcement, it facilitates discrimination learning. Conversely, a response-contingent CS previously associated with the absence of shock (AvCS?) retards discrimination learning. To evaluate whether these findings reflect across-reinforcement blocking and enhancement effects, two experiments investigated the effects of appetitively conditioned stimuli on fear conditioning to a novel stimulus that was serially compounded with the appetitive CS during conditioned-emotional-response (CER) training. Although there were no differential effects of the appetitive CSs in CER acquisition, Experiment 1, using a relatively weak shock US, showed that a CS previously associated with food (ApCS+) retarded CER extinction to the novel stimulus, in evidence of enhanced fear conditioning to that stimulus. In addition, Experiment 2, using a stronger shock US, showed that a CS previously associated with the absence of food (ApCS?) facilitated CER extinction to the novel stimulus, in evidence of weaker fear conditioning to that stimulus. These results parallel traditional blocking effects and indicate not only that an ApCS+ and an ApCS? are functionally similar to AvCSs of opposite sign, but that their functional similarity is mediated by common central emotional states.     

Recovery from conditioned suppression to a partially overlapping compound stimulus

In a conditioned suppression paradigm, a partially overlapping compound stimulus signaled occurrences of electric shock. That compound CS consisted of 3 min of continuous illumination of the houselight with three discrete 5-see presentations of a tone superimposed. A .5-sec electric shock was coterminous with each tone presentation. Rats that received this treatment in early conditioning sessions showed considerable recovery from conditioned suppression to the houselight as the experiment progressed. However, the effect was not reversible, and it could not be demonstrated in rats that experienced extensive prior conditioning to the houselight alone. These results are discussed in relation to a hypothesis concerning the modulation of behavioral control exerted by elements of compound stimuli and as they relate to a recent theoretical model for Pavlovian fear conditioning.     

Pavlovian conditioning in multiple contexts: Competition between contexts for comparator status

Water-deprived rats were used to investigate the effects of training a CS in more than one context on conditioned lick suppression. In each experiment, partial reinforcement of the CS was intermingled with unsignaled presentations of the US. In Experiment 1, subjects were either trained in one context alone, trained consecutively in two contexts (such that all training in one context occurred prior to any training in the second context), or trained alternately in two contexts. Following training, the first context, the second context, or neither context was extinguished. Testing of the CS occurred in a third (neutral) context. To the extent that either training context became established as a comparator stimulus for the CS, the comparator hypothesis (Miller & Matzel, 1988) predicts an increase in excitatory responding to the CS following extinction of that context. Subjects trained in a single context exhibited appreciable fear of the CS only when the CS’s training context had been extinguished. Additionally, subjects trained consecutively in the two contexts showed increased fear of the CS following extinction of the second, but not the first training context (i.e., a recency effect). Subjects trained alternately in the two contexts showed no increased fear of the CS as a result of either context alone being extinguished. In Experiment 2, subjects trained alternately in two contexts showed increased fear of the CS only when both training contexts were extinguished, suggesting that both training contexts had become comparator stimuli. These data indicate that multiple training contexts can either compete or act synergistic-ally in modulating responding to a Pavlovian trained CS as a function of the order of training in the different contexts.     

Order and duration of stimuli are important determinants of reactivation

Prior cuing treatments intended to alleviate the forgetting of a conditioned avexsion-to an odor were tested with 18-day-old rats. Previous experiments had shown that when such pups were conditioned with the use of a CS?/CS+ procedure, pretest presentation of the CS? or US, but not the CS+, alleviated the forgetting otherwise seen after a 3-h retention interval. In Experiment 1, it was determined that the forgetting was not alleviated if the GS? was either preceded or followed by presentation of the CS+, despite the fact that the CS?/CS+ ordering mimicked that of original conditioning. Experiment 2 was an examination of the balance of extinction and reactivation effects caused by presenting the CS+ for varying durations following the 3-h retention interval. The forgetting over this interval was alleviated if the CS+ was presented for 5 or 15 sec, but not 30 sec. With an increase in duration of exposure from 15 to 30 sec, the consequences of the CS+ as a prior cuing treatment apparently shifted from reactivation to extinction. Experiment 3 was a test of the interaction between the consequences of different lengths of CS+ exposure and the effectiveness of adding CS? to the CS+ as a reactivation treatment. The varied effectiveness of reactivation treatments is discussed interms of a change in stimulus conditions from training to reactivation.