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.     

Transfer of negative occasion setting and conditioned inhibition across conditioned and unconditioned stimuli

The transfer of negative occasion setting and conditioned inhibition across conditioned stimuli (CSs) and unconditioned stimuli (USs) was examined in four experiments that used Pavlovian appetitive feature negative discrimination training procedures with rats. After training with simultaneous compounds (A+, XA-), X inhibited conditioned responding (CRs) elicited by other CSs and CRs supported by other appetitive USs that had not been involved in discrimination training. After training with serial compounds (A+, X----A-), X's power to set the occasion for nonresponding transferred across CSs and USs only if those events had also been involved in serial feature negative discrimination training. The results supported the suggestion that the acquisition of negative occasion setting involves the representation of individual events in a higher order memory system, separate from that involved in simple association, and that negative occasion setters act only on events that are represented in that system.     

Effect of telencephalon ablation on the reinforcing and eliciting properties of species-specific events in Betta splendens

In male Betta splendens, aggressive behavior is drastically attenuated following telencephalon ablation. Because instrumental training and Pavlovian conditioning experiments with intact fish have suggested that associative factors may play an important role in the performance of agonistic behaviors, the effect of ablation on instrumental learning and Pavlovian conditioning was studied. In Experiment 1, ablation had no effect on the learning of the instrumental tunnel-swimming response reinforced by mirror presentation (i.e., viewing a conspecific), although the mirror presentations in yoked-control groups elicited fewer responses in ablates than in normal and sham-operated control fish. Yoked controls further established that instrumental responding was maintained by the reinforcement contingency and was not merely the result of increased motor activity. Experiment 2 studied Pavlovian conditioning of the components of the agonistic display. Unconditioned fin erection, gill erection, and tail beating (i.e., unconditioned responses, URs) to the mirror US all were less frequent in ablates than in normals or shams. Of these, only gill cover erection showed evidence of true conditioning (i.e., conditioned responses; CRs) in which responses to the conditioned stimulus (CS) are due to the pairings of CS and US (unconditioned stimulus). However, ablates suffered no impairment of conditioned gill erections. Ablates performed fewer fin erections to the CS; however, fin erection responses were not due to CS-US pairings but were attributable to pseudoconditioning. These results are related to hypotheses postulating the involvement of learning mechanisms in ablation-produced deficits and normal aggressive behavior.     

Effects of long- and variable-duration signals for food on activity,instrumental responding,and eating

Incentive-motivation theories typically assume that the conditioning of appetitive motivation involves the same parameters as Pavlovian salivary conditioning. In contrast, the Soltysik-Konorski model asserts that drive is inhibited by stimuli closely associated with food (salivary CSs) and augmented by stimuli more loosely associated with food (long and variable CS-US interval). Experiment 1 examined this latter proposition. Sixty-four rats were given extensive exposure to each of four environmental CSs, two while hungry and two while satiated. Within each deprivation condition, food was given 30–300 sec after placement of the rats in one environment, and was not given in the other environment. Performance on three separate measures—activity, lever-pressing, and food consumption—was higher in the environments previously associated with food. Experiment 2 examined the effects of discrete stimuli presented in advance of eating; in accord with the results of Experiment 1, food consumption was greater after a stimulus (1- to 9-min duration) previously paired with food than after no stimulus or after a stimulus unpaired with food. The overall results indicate (a) that stimuli associated with food become capable of facilitating a variety of food-directed behaviors, possibly via the conditioning of a common appetitive system, and (b) that a close association between the stimuli and food is not essential for such conditing to occur.     

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.     

Conditioned inhibition produced by extinction-mediated recovery from the relative stimulus validity effect: a test of acquisition and performance models of empirical retrospective revaluation

Empirical retrospective revaluation is a phenomenon of Pavlovian conditioning and human causal judgment in which posttraining changes in the conditioned response (Pavlovian task) or causal rating (causal judgment task) of a cue occurs in the absence of further training with that cue. Two experiments tested the contrasting predictions made by 2 families of models concerning retrospective revaluation effects. In a conditioned lick-suppression task, rats were given relative stimulus validity training, consisting of reinforcing a compound of conditioned stimuli (CSs) A and X and nonreinforcement of a compound of CSs B and X, which resulted in low conditioned responding to CS X. Massive posttraining extinction of CS A not only enhanced excitatory responding to CS X, but caused CS B to pass both summation (Experiment 1) and retardation (Experiment 2) tests for conditioned inhibition. The inhibitory status of CS B is predicted by the performance-focused extended comparator hypothesis (J. C. Denniston, H. I. Savastano, & R. R. Miller, 2001), but not by acquisition-focused models of empirical retrospective revaluation (e.g., A. Dickinson & J. Burke, 1996; L. J. Van Hamme & E. A. Wasserman, 1994).     

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.     

Aversive CS control of instrumental avoidance as a function of selected parameters and method of Pavlovian conditioning

Effects of aversive CSs upon dogs' instrumental avoidance responding were assessed as a function of the Pavlovian conditioning parameters using off-baseline (Experiment 1) and on-baseline (Experiment 2) conditioning procedures. In each experiment, three UCS intensitites (2, 6, and 10 mA) were crossed factorially with two CS-UCS intervals (10 and 90 sec) yielding six groups. After 4 days of conditioning, the CS-produced effects on avoidance were determined primarily by an interaction between Pavlovian procedure (off- versus on-baseline) and UCS intensity. With off-baseline conditioning, CS-produced facilitation of avoidance was an increasing function of the UCS intensity. However, with on-baseline conditioning, only the CS paired with weak UCS facilitated avoidance, and the CS paired with the strong UCS suppressed responding. This reversing parametric function poses problems for two-process motivational theories of avoidance.     

Pavlovian conditioning of sexual arousal: parametric and background manipulations

Recent studies have shown that a Pavlovian conditioned stimulus (CS) for unconsummated sexual arousal can increase the rate of copulation in the rat. This report includes five experiments examining the effects of parametric manipulations on the conditioned arousal response. Results show that between six and nine trials are necessary for reliable conditioning, but extinction is somewhat slower than acquisition. The function for the CS-US (unconditioned stimulus) interval is quadratic, with a minimum of several minutes required for effective conditioning. In the first three experiments, it appeared that background cues were conditioned as well as the designated CSs, and this was tested explicitly in the last two studies. In one, the effect of background cues was shown by training and testing in different situations; in the second, background cues were shown to be subject to latent inhibition. These results demonstrate the influence of Pavlovian learning in sexual behavior and help to provide the basis for an animal model of the acquisition of deviant sexual arousal in humans.     

Effects of contextual conditioning and unconditional stimulus presentation on performance in appetitive conditioning

Four experiments with rat subjects examined the effects of contextual conditioning on conditioned appetitive performance. Experiment 1 compared the effects of contextual conditioning on performance to conditioned stimuli (CSs) with different conditioning histories. Contextual conditioning enhanced performance to the CS if the CS had first been conditioned and then extinguished, but had no effect on performance when the CS had been merely paired or unpaired with food. Experiments 2 and 3 then asked whether the effect on the extinguished CS was due to contextual conditioning acting as a cue for conditioning. In Experiment 2, extinction procedures in which extra unconditioned stimuli (USs) were presented during the intertrial intervals were found to reduce the CS's sensitivity to enhancement by contextual conditioning, but had no effect on spontaneous recovery. In Experiment 3, USs added to conditioning or extinction acquired the ability to cue the corresponding performance. Under some conditions, USs added to conditioning could suppress performance (Experiment 4). The results suggest that contextual conditioning has complex effects that can be better understood by recognizing that contextual conditioning, as well as the USs that create it,Mayacquire discriminative control over conditioned responding.     

Blocking in successive differential conditioning: Prior acquisition of control by internal cues blocks the acquisition of control by brightness

In each of four experiments the acquisition of discriminative stimulus control by brightness cues (black vs white runway) in a successive go/no-go instrumental discrimination was blocked in groups given prior discrimination training with internal (reward-produced and intertrial interval-related) cues as relevant discriminanda and brightness cues irrelevant. The blocking effects obtained here in instrumental conditioning were substantial and in most cases complete. Blocking occurred whether brightness in Phase I varied across trials but was uncorrelated with reinforcement (varied-irrelevant, V-I, condition) or whether all Phase 1 trials occurred to a single value on the brightness dimension (constant-irrelevant, C-I, condition) which then served as a redundant S+ cue in Phase 2 while the previously unexperienced brightness cue was added to the S- stimulus compound, or vice versa. Blocking in the V-I condition was shown not to be due simply to nondifferential reinforcement of brightness in Phase 1, but to depend on the prior acquisition of discriminative stimulus control by internal cues. As in Pavlovian conditioning, blocking here was an increasing function of amount of prior conditioning to the blocking stimulus. The results encourage the prospect that the procedures used here can be developed into a viable instrumental conditioning companion to the Pavlovian procedures now used almost exclusively to study blocking.     

Posttraining prefrontal lesions impair jaw movement conditioning performance,but have no effect on accompanying heart rate changes

This experiment examined the role of the medial prefrontal cortex (mPFC) in regulating learned autonomic and somatomotor responses in rabbits using appetitive Pavlovian conditioning. Interstimulus interval (ISI) duration [i.e., the time between the onset of the conditioned stimulus (CS) and unconditioned stimulus (US)] was manipulated in order to determine whether ISI duration was related to the heart rate (HR) responses obtained during conditioning. Two groups received either a 1- or a 4-s ISI, with a tone as the CS and an intraoral pulse of water as the US. Another two groups received explicitly unpaired presentations of either the 1- or 4-s tone CS and water US. Few conditioned jaw movement (JM) or HR conditioned responses (CRs) were observed in the unpaired conditions. Significant JM conditioning was, however, elicited by the paired conditions, especially to the 4-s ISI. Consistent CS-evoked HR accelerations were observed in both ISI conditions. After five sessions of training, the mPFC was lesioned in half the animals. A separate group of paired animals received sham lesions. After surgical recovery, all animals received 3 days of postoperative training. During the first postoperative training session, JM CRs significantly declined in both groups with mPFC lesions in comparison to the groups with sham lesions. The mPFC lesions, however, did not affect the CS-evoked cardiac accelerations, which again occurred during postoperative training.     

Are anticipatory first and second interval skin conductance responses indicators of predicted aversiveness?

The experiment reviewed here was an attempt to examine whether or not magnitudes of first and second interval skin conductance responses (FIR, SIR) to different conditioned stimuli (CS), each predicting another unconditioned stimulus (US), mirror the aversiveness of the predicted USs. Twenty human subjects received a differential Pavlovian conditioning training with five CSs, each one paired eight times with a specific US. Magnitudes of FIR and SIR elicited by CSs increased with increasing aversiveness of the USs predicted. It is concluded therefore that the SIR is not a mere indicator of a cognitive expectancy process but is also affected by emotional aspects.     

An analysis of sequential variables in Pavlovian conditioning employing extended and limited acquisition training

Sequential theory’s memory model of learning has been successfully applied in response contingent instrumental conditioning experiments (Capaldi, 1966, Capaldi, 1967, Capaldi, 1994 and Capaldi and Miller, 2003). However, it has not been systematically tested in nonresponse contingent Pavlovian conditioning experiments. The present experiments attempted to determine if several sequential variables affect responding in Pavlovian situations as they do in instrumental ones. Of primary concern here were the effects on extinction of number of NR transitions (the number of times a nonreinforced trial is followed by a reinforced trial), N-length (the number of successive nonreinforced trials that precede a reinforced trial), and percentage of reinforcement (50 versus 100%) following either extended acquisition training (Experiment 1, 720 trials) or limited acquisition training (Experiment 3, 24 trials). In agreement with a sequential analysis, N-length increased resistance to extinction more than number of NR transitions following extensive training with the opposite occurring following limited training. In Experiment 1, greater resistance to extinction was associated with 50% than with 100% reinforcement, a partial reinforcement extinction effect (PREE). Experiment 2 examined an anomalous finding obtained in Experiment 1. A major theoretical difference between instrumental and Pavlovian conditioning has been held to be the greater ease of producing a PREE in instrumental than in Pavlovian conditioning (Kimble, 1961 and Mackintosh, 1974). However, the findings obtained here suggest that the probability of obtaining a PREE and other Pavlovian extinction effects, as in instrumental conditioning, increases along with the effectiveness of the sequential variables employed.     

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.     

The pavlovian analysis of instrumental conditioning

An account was given of the development within the Russian literature of a uniprocess formulation of classical and instrumental conditioning, known as the bidirectional conditioning hypothesis. The hypothesis purports to offer a single set of Pavlovian principles to account for both paradigms, based upon a neural model which assumes that bidirectional (forward and backward) connections are formed in both classical and instrumental conditioning situations. In instrumental conditioning, the bidirectional connections are hypothesized to be simply more complex than those in classical conditioning, and any differences in empirical functions are presumed to lie not in difference in mechanism, but in the strength of the forward and backward connections. Although bidirectional connections are assumed to develop in instrumental conditioning, the experimental investigation of the bidirectional conditioning hypothesis has been essentially restricted to the classical conditioning operations of pairing two CSs (sensory preconditioning training), a US followed by a CS (backward conditioning training) and two USs. However, the paradigm involving the pairing of two USs, because of theoretical and analytical considerations, is the one most commonly employed by Russian investigators. The results of an initial experiment involving the pairing of two USs, and reference to the results of a more extensive investigation, leads us to tentatively question the validity of the bidirectional conditioning account of instrumental conditioning.     

Comparison of Single Unit Responses to Tone, Light, and Compound Conditioned Stimuli during Rabbit Classical Eyeblink Conditioning

Unit recordings and lesion studies have implicated the cerebellum as an essential site for the acquisition and maintenance of the conditioned eyeblink response. The current study looked at the neural characteristics of conditioned stimulus (CS) processing in the interpositus nucleus of the cerebellum after training New Zealand white rabbits (Oryctolagus cuniculus) in one of two conditioning paradigms: (a) compound conditioning (CMP), a compound CS consisting of light and tone paired with an air puff unconditioned stimulus (US); or (b) stimulus compounding (ALT), alternating blocks of tone CS and light CS trials paired with the air puff US. Single unit responses were recorded during five sessions after the animals had reached an asymptotic level of responding. Animals were tested for behavioral and neural responses to CS alone trials that included tone alone, light alone, and compound tone-light trials. For the CMP group, the compound CS elicited 80 to 90% conditioned eyeblink responses (CRs), whereas the individual tone and light CSs elicited only 40 to 50% CRs. For the ALT group, all three CSs (tone, light, and compound) elicited very high levels of responding of at least 80% CRs. For the CMP group, there were roughly equal numbers of cells responding to all of the CSs. This includes cells that responded exclusively to one, and only one, of the three stimuli and also those cells that responded to combinations of two or more. Cells from the ALT group were far more likely to respond exclusively to only one of the CSs. Both the behavioral and physiological results suggest that the compound tone-light stimulus was processed as a distinct stimulus, separate from the component tone and light. These results are discussed in the context of multisensory processing.     

Pavlovian heart rate and jaw movement conditioning in the rabbit: effects of medial prefrontal lesions

An experiment was conducted in which jaw movements (JM) and heart rate (HR) were concomitantly assessed in rabbits during simple Pavlovian conditioning. A 2-s 1200-Hz tone was the conditioned stimulus (CS) and an intraoral 1-cc pulse of 0.5 M sucrose-water solution was the unconditioned stimulus (US). Sham and medial prefrontal (mPFC)-lesioned animals received paired CS/US training with a 70- to 75-dB CS and were compared with sham- and mPFC-lesioned animals that received explicitly unpaired CS/US presentations. The percentages of JM CRs were significantly greater in the paired than the unpaired groups, but mPFC lesions had no effect on this measure. Conditioned HR decelerations occurred only in the paired groups and then only during the first session of training. Moreover, these CS-evoked cardiac decelerations were somewhat attenuated by the mPFC lesion. CS-evoked HR accelerations, which were significantly greater in unpaired than in paired animals, occurred during the four subsequent sessions. These results suggest that a CS-evoked cardioinhibitory process, mediated by the mPFC, is engendered by Pavlovian appetitive conditioning, as has been previously demonstrated for aversive conditioning. However, during JM conditioning these inhibitory changes are quickly replaced by tachycardia, possibly related to increased nonspecific somatomotor activity, since the tachycardia was somewhat greater in the unpaired animals.     

Contingency theory and the effect of the duration of signals for noncontingent reinforcement

Rats learned to approach a light that signaled food when food occurred only after the signal, but did not approach the light when food was equally probable regardless of the signal. When food unpaired with the light was preceded by a 5-sec noise, little learning was evident as well, but when the unpaired food was preceded by a 15-sec noise, approach to the light conditioned stimulus readily occurred. These effects of signal duration are similar to those previously obtained with the effect of free food on free-operant behavior and challenge existing theories of contingency effects in Pavlovian conditioning.