Differential effects of forward or simultaneous conditioned stimulus-unconditioned stimulus intervals on the defensive behavior system of the Norway rat (Rattus norvegicus)

To test several predictions derived from a behavior-systems approach, the authors assessed Pavlovian fear conditioning in rats after 30 trials of forward, simultaneous, or unpaired training. Direct evidence of conditioned fear was collected through observation of flight and freezing reactions during presentations of the conditioned stimulus (CS) alone. The authors also tested the CS's potential to reinforce an instrumental escape response in an escape-from-fear paradigm. On the one hand, rats that received forward training showed conditioned freezing, but no conditioned flight was observed. On the other hand, rats that received simultaneous training showed conditioned flight, but no conditioned freezing was observed. Rats that received either forward or simultaneous pairings showed instrumental learning of the escape-from-fear response. Implications for several theories of Pavlovian conditioning are discussed.     

Pavlovian conditioning in human skilled motor behavior

The role of Pavlovian contingencies in human skilled motor behavior was investigated in three experiments by means of a new conditioning preparation. In Experiment 1 the present method was shown to be appropriate for the study of associative learning. Subjects who experienced a standard delay configuration performed significantly more conditioned responses than subjects who received either backward conditioning or random pairings. Stimulus generalization was shown to be slight in two additional groups. Subsequent experiments examined conditioning with multiple conditioned stimuli (CSs). In particular, in Experiment 2 some reciprocal overshadowing was demonstrated when two conditional stimuli (tone and vibration) were compounded. Experiment 3 investigated blocking. Blocking was less than expected, however. Subjects’ perceptions of the stimuli and reaction time data suggest that a certain proportion had shifted their attention to the added element of the CS compound. Results are discussed in relation to other studies on Pavlovian learning in humans and animals, which are concerned with “stimulus selection.”     

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.     

Concomitant Pavlovian conditioning of heart rate and leg flexion responses in the rat

Pavlovian conditioning was studied in male Fischer 344 rats using tones as the conditioned stimulus (CS) and footshock as the unconditioned stimulus (UCS). Different groups of animals received (a) contiguous CS-UCS pairings with a 0.5 sec CS, (b) contiguous CS-UCS pairings with a 4.0 sec CS, or (c) a random sequence of noncontiguous tones and shocks using either a 0.5 sec or a 4.0 sec CS. Heart rate (HR) and leg flexion (LF) responses were recorded. Leg flexion conditioning occurred only in the 0.5 sec contiguous group. Decelerative HR CRs occurred only in the 4.0 sec contiguous group. Accelerative HR changes occurred in the other two groups but were significantly greater in the 0.5 sec contiguous group. These results are similar to but not identical to those obtained during eyeblink or nictitating membrane conditioning in rabbits, and suggest that the topography of the Pavlovian HR CR is dependent on the simultaneous occurrence of other classically conditioned responses.     

Contextual control of inhibition with reinforcement: adaptation and timing mechanisms

Four experiments with rats studied the effects of switching the context after Pavlovian conditioning. In three conditioned suppression experiments, a large number of conditioning trials created "inhibition with reinforcement" (IWR), in which fear of the conditional stimulus (CS) reached a maximum and then declined despite continued CS-unconditional stimulus pairings. When IWR occurred, a context switch augmented fear of the CS; IWR and augmentation were highly correlated. Neither IWR nor augmentation resulted from inhibition of delay (IOD): In conditioned suppression, IWR and augmentation occurred without IOD (Experiment 3), and in appetitive conditioning (Experiment 4), IOD occurred without IWR or augmentation. IWR may occur in conditioned suppression because the animal adapts to fear of the CS in a context-specific manner. The authors discuss several implications.     

Pavlovian second-order conditioned analgesia

Three experiments with rat subjects assessed conditioned analgesia in a Pavlovian second-order conditioning procedure by using inhibition of responding to thermal stimulation as an index of pain sensitivity. In Experiment 1, rats receiving second-order conditioning showed longer response latencies during a test of pain sensitivity in the presence of the second-order conditioned stimulus (CS) than rats receiving appropriate control procedures. Experiment 2 found that extinction of the first-order CS had no effect on established second-order conditioned analgesia. Experiment 3 evaluated the effects of post second-order conditioning pairings of morphine and the shock unconditioned stimulus (US). Rats receiving paired morphine-shock presentations showed significantly shorter response latencies during a hot-plate test of pain sensitivity in the presence of the second-order CS than did groups of rats receiving various control procedures; second-order analgesia was attenuated. These data extend the associative account of conditioned analgesia to second-order conditioning situations and are discussed in terms of the mediation of both first- and second-order analgesia by an association between the CS and a representation or expectancy of the US, which may directly activate endogenous pain inhibition systems.     

Backward conditioning: mediation by the context

The information acquired in backward conditioning (i.e., outcome-->cue) was assessed in 3 Pavlovian lick-suppression experiments with water-deprived rats as subjects. Experiment 1 confirmed previous research that few outcome-->cue pairings made the cue into a conditioned excitor and additionally showed that massive posttraining extinction of the training context attenuated a backward-trained cue's excitatory value. Experiment 2 found that many outcome-->cue pairings made the cue into a conditioned inhibitor and that the same context manipulation attenuated this inhibitory value. Experiment 3 confirmed the observations of Experiments 1 and 2 and demonstrated that these effects of context extinction were specific to backward-trained cues conditioned in the extinguished context. These results are interpreted in terms of cue-->context and context-->outcome associations.     

Instrumental to Pavlovian transfer: Learning about response-reinforcer contingencies affects subsequent learning about stimulus-reinforcer contingencies

Two experiments examined the interaction between response-reinforcer (R-S) and stimulus-reinforcer (S-S) learning. In both experiments, three groups of rats were exposed to escapable, yoked inescapable, or no shocks. All groups were then exposed to either two or four sessions of truly random control (TRC) conditioning (Experiment 1) or to an excitatory conditioning procedure (Experiment 2) in which the shock US occurred with either moderate or low probability. Excitatory strength of the CS was assessed during extinction by a conditioned emotional response (CER) test. Inescapably shocked rats conditioned less than did their escapably shocked and nonshocked partners under all TRC conditions of Experiment 1, but only conditioned less than their partners in Experiment 2 when exposed to the moderate CS-US contingency for four sessions. These results provide a clear demonstration of transfer between instrumental training and Pavlovian excitatory conditioning and thus, support the influence of learning about R-S contingencies upon subsequent learning about S-S contingencies. Both a contextual blocking interpretation and an expansion of the learned helplessness theory were discussed as possible explanations of this transfer.     

Instrumental learning in hyperdopaminergic mice

In two experiments we investigated the effects of elevated dopaminergic tone on instrumental learning and performance using dopamine transporter knockdown (DAT KD) mice. In Experiment 1, we showed that both DAT KD mice and wild-type controls were similarly sensitive to outcome devaluation induced by sensory specific satiety, indicating normal action-outcome learning in both groups. In Experiment 2, we used a Pavlovian-to-instrumental transfer procedure to assess the potentiation of instrumental responding by Pavlovian conditional stimuli (CS). Although during the Pavlovian training phase the DAT KD mice entered the food magazine more frequently in the absence of the CS, when tested later both groups showed outcome-selective PIT. These results suggest that the elevated dopaminergic tone reduced the selectivity of stimulus control over conditioned behavior, but did not affect instrumental learning.     

Blocking and sensory preconditioning effects in morphine analgesic tolerance: Support for a pavlovian conditioning model of drug tolerance

Two experiments on rats tested predictions of a Pavlovian conditioning model of drug tolerance which holds that tolerance is the result of compensatory conditioned responses, developed to environmental stimuli accompanying the drug administrations, which attenuate the direct effects of the drug. In Experiment I, the acquisition of tolerance-modulating properties by the tone component of a tone-light compound stimulus which accompanied morphine administrations was reduced by prior light-morphine pairings (blocking). In Experiment II, a tone stimulus acquired tolerance-modulating properties through prior pairings with a light stimulus which later accompanied morphine administrations (sensory preconditioning). These findings are uniquely predicted by the Pavlovian conditioning model of drug tolerance and are incompatible with traditional theories which assign no role to environmental stimuli present at the time of drug administration.     

Blocking and unblocking of conditioned analgesia

Two experiments with rat subjects assessed the blocking phenomenon using inhibition of responding to painful thermal stimulation as an index of conditioned analgesia established through conditioned stimulus (CS)-shock pairings. In Experiment 1, a group of rats receiving the standard blocking procedure ($\text{A+}\text{AB+}$) showed shorter response latencies during a hot plate test of pain sensitivity in the presence of the added CS than groups of rats receiving various blocking control procedures. Experiment 2 replicated the blocking outcome of Experiment 1 and also demonstrated that the addition of a second shock unconditioned stimulus (US) during the compound conditioning phase attenuated the blocking effect; i.e., unblocking was observed. However, the deletion of a second shock US during compound conditioning in a group that had received two shocks on each trial in the first phase of the experiment failed to result in unblocking. These data extend the associative account of conditioned analgesia to situations involving stimulus selection and the implications for the analysis of aversive learning are discussed.     

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.     

Degraded contingency revisited: posttraining extinction of a cover stimulus attenuates a target cue's behavioral control

In a Pavlovian conditioning situation, unsignaled outcome presentations interspersed among cue-outcome pairings attenuate conditioned responding to the cue (i.e., the degraded contingency effect). However, if a nontarget cue signals these added outcomes, responding to the target cue is partially restored (i.e., the cover stimulus effect). In 2 conditioned suppression experiments using rats, the effect of posttraining extinction of the cover stimulus was examined. Experiment 1 found that this treatment yielded reduced responding to the target cue. Experiment 2 replicated this finding, while demonstrating that this basic effect was not due to acquired equivalence between the target cue and the cover stimulus. These results are consistent with the extended comparator hypothesis interpretation of the degraded contingency and cover stimulus effects.     

Pavlovian conditioning of Hermissenda: current cellular, molecular, and circuit perspectives

The less-complex central nervous system of many invertebrates make them attractive for not only the molecular analysis of the associative learning and memory, but also in determining how neural circuits are modified by learning to generate changes in behavior. The nudibranch mollusk Hermissenda crassicornis is a preparation that has contributed to an understanding of cellular and molecular mechanisms of Pavlovian conditioning. Identified neurons in the conditioned stimulus (CS) pathway have been studied in detail using biophysical, biochemical, and molecular techniques. These studies have resulted in the identification and characterization of specific membrane conductances contributing to enhanced excitability and synaptic facilitation in the CS pathway of conditioned animals. Second-messenger systems activated by the CS and US have been examined, and proteins that are regulated by one-trial and multi-trial Pavlovian conditioning have been identified in the CS pathway. The recent progress that has been made in the identification of the neural circuitry supporting the unconditioned response (UR) and conditioned response (CR) now provides for the opportunity to understand how Pavlovian conditioning is expressed in behavior.     

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.     

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.     

Asymmetrical effects of Pavlovian excitatory and inhibitory aversive transfer on Pavlovian appetitive responding and acquisition

Predictions of a theory of Pavlovian motivational transfer, which incorporates principles of both the theory of reciprocal inhibition and the Rescorla-Wagner model, were tested in several Pavlovian aversive to Pavlovian appetitive transfer tasks. As predicted, the presence of a signal for an aversive event, conditioned stimulus (AV CS+), reliably suppressed performance of appetitive conditioned responses (CRs) whether imposed during acquisition or on independently established responding. Acquisition of appetitive responding to a novel CS reinforced in compound with an AV CS+, however, was enhanced (“superconditioning”). This observation suggests that the effects of a discrepancy between expectation and actual outcome on a conditioning trial are influenced by the affective value of both the expectation and the reinforcer. These transfer effects were not symmetrical for an inhibitory aversive stimulus (AV CS?). An AV CS? did not enhance appetitive responding compared to a random control condition, nor did the AV CS? reduce (i.e., block) appetitive conditioning to a novel CS when appetitive reinforcement occurred in the presence of the AV CS?. Comparison of the two shock-exposed conditions with a naive control condition suggests that previous results that were apparently consistent with inhibitory aversive enhancement and blocking of appetitive conditioning may have been due to aversive context conditioning.     

Backward conditioning as an extinction procedure

In two conditioned suppression experiments, rats received Pavlovian forward defense conditioning in which tonal conditioned stimuli (CSs) terminated with the onset of scrambled grid shock unconditioned stimuli (USs). After this experience, the rats then received a Pavlovian backward conditioning procedure in which the same USs now terminated with the onset of the same CSs. Although the two experiments differed greatly in terms of CS and US parameters, number of forward and backward pairings, and in terms of the general techniques used to establish and measure the Pavlovian conditioned response (CR), the results of both experiments agreed in showing that backward conditioning can indeed weaken a CR based on forward pairings. The results also show that, under some conditions, the backward procedure can be at least as effective in weakening an established CR as the traditional CS-alone extinction procedure; but, under other conditions, the backward procedure is less effective and leads to more spontaneous recovery than the CS-alone procedure.     

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.     

Pavlovian to instrumental transfer of control in a human learning task

Pavlovian learning tasks have been widely used as tools to understand basic cognitive and emotional processes in humans. The present studies investigated one particular task, Pavlovian-to-instrumental transfer (PIT), with human participants in an effort to examine potential cognitive and emotional effects of Pavlovian cues upon instrumentally trained performance. In two experiments, subjects first learned two separate instrumental response-outcome relationships (i.e., R1-O1 and R2-O2) and then were exposed to various stimulus-outcome relationships (i.e., S1-O1, S2-O2, S3-O3, and S4-) before the effects of the Pavlovian stimuli on instrumental responding were assessed during a non-reinforced test. In Experiment 1, instrumental responding was established using a positive-reinforcement procedure, whereas in Experiment 2, a quasi-avoidance learning task was used. In both cases, the Pavlovian stimuli exerted selective control over instrumental responding, whereby S1 and S2 selectively elevated the instrumental response with which it shared an outcome. In addition, in Experiment 2, S3 exerted a nonselective transfer of control effect, whereby both responses were elevated over baseline levels. These data identify two ways, one specific and one general, in which Pavlovian processes can exert control over instrumental responding in human learning paradigms, suggesting that this method may serve as a useful tool in the study of basic cognitive and emotional processes in human learning.