Effects of US parameters on classical conditioning of cat hindlimb flexion

Unconditioned stimulus (US) intensity and duration were manipulated to determine their effects on cat hindlimb flexion conditioning. Seven consecutive days of acquisition training of a hindlimb flexor response to a tone conditioned stimulus (CS) were followed by 2 days of extinction. Eight animals in each of 12 groups received a 1-, 2-, 3-, or 4-mA, 60-Hz shock delivered to the right hindleg for 25, 50, or 100 msec as the US. Analysis of conditioned-response (CR) frequency indicated that conditioned responding was a positive function of both the intensity and duration of shock, although these variables did not interact with one another. CR latency and amplitude were decreased and increased, respectively, by increases in US intensity. The pattern of results reported here may support a contiguity notion of conditioning, and are discussed in the context of other conditioning preparations.     

Evaluating the TD model of classical conditioning

The temporal-difference (TD) algorithm from reinforcement learning provides a simple method for incrementally learning predictions of upcoming events. Applied to classical conditioning, TD models suppose that animals learn a real-time prediction of the unconditioned stimulus (US) on the basis of all available conditioned stimuli (CSs). In the TD model, similar to other error-correction models, learning is driven by prediction errors-the difference between the change in US prediction and the actual US. With the TD model, however, learning occurs continuously from moment to moment and is not artificially constrained to occur in trials. Accordingly, a key feature of any TD model is the assumption about the representation of a CS on a moment-to-moment basis. Here, we evaluate the performance of the TD model with a heretofore unexplored range of classical conditioning tasks. To do so, we consider three stimulus representations that vary in their degree of temporal generalization and evaluate how the representation influences the performance of the TD model on these conditioning tasks.     

Stepwise reductions in US density and intensity: Maintenance and extinction of conditioned suppression

Information was sought on how the strength of fear, acquired and extinguished in a classical conditioning paradigm, might be affected by certain of the circumstances that normally are associated with the course of avoidance learning. Following preconditioning exposures to the conditioned stimulus, one group of rats (Group PRF) was given continuous conditioned stimulus-unconditioned stimulus (CS-US) acquisition trials followed by stepwise reductions in US probability (given the CS) over three phases to about 11% for the final phase. Another group, Group US(lo), was given continuous CS-US pairings throughout, but, following acquisition, received stepwise reductions in US intensity (to permit evaluation of a progressively changing feature of the US over a wide range without having to employ excessive shock) over the same phases. A third group, Group US(hi), received unchanging (from acquisition) CS-US pairings over these phases, and an explicitly unpaired control group (Group RU) was included. Although outcomes differed somewhat depending upon whether suppression ratios or absolute measures of responding were considered, the major findings were that suppression to the CS was complete by the end of acquisition and persisted thereafter throughout the three phases for all but control subjects. In contrast to Group US(lo), which showed relatively little resistance to extinction of suppression to subsequent CS-only exposures, Groups US(hi) and PRF displayed marked resistance over two separate, extended sets of extinction sessions. Suppression to context cues was pronounced only for Groups US(hi) and RU, and varied as a function of US parameters and not whether the shocks were or were not signaled. Theoretical reconciliation of these findings was most difficult for Groun PRF.     

Motivational vs. associative role of the US in classical conditioning of the rabbit’s nictitating membrane response

Two experiments tested the motivational role of the US in classical conditioning of the rabbit’s nictitating membrane (NM) response. In Experiment 1, subjects were trained to an intermediate performance level and then given a series of (1) CS-US trials, (2) “backwards” US-CS trials, (3) CS-alone trials, (4) US-alone trials, or (5) no-stimulus presentations. Interpolated presentations of the US, either alone or in a backwards contingency, tended to produce an impairment of subsequent acquisition. In Experiment 2, subjects were trained with strong or weak US intensity on paired or interpolated trials. US intensity on interpolated trials had only a very small effect, whereas the effect of US intensity on paired trials was quite large. Shifts in paired-trial US intensity produced corresponding shifts in performance, but shifts in the intensity of the interpolated US produced no apparent effect. We conclude that the arousal of motivation is not sufficient to maintain performance in classical NM conditioning.     

Defensive responding and leverpress topography compatibility: Effects of shock intensity,S-S interval,and lever position

The degree of compatibility between defensive responding and leverpress topography was investigated. Rats were trained to leverpress on a Sidman avoidance schedule for one 60-min session. Measures of defensive responding (jumping and rearing) were recorded during initial training. Shock intensity (.5, 1.0, and 2.0 mA) and S-S interval (1, 3, and 5 sec) were varied in Experiment 1. S-S interval had no effect on defensive responding or on performance. Rate of jumping was an increasing function of intensity only at lower intensities. Both jumping and rearing were significantly correlated with leverpressing. Shock intensity (1.0 and 2.0 mA) and lever position (3.02 and 16.83 cm) were varied in Experiment 2. Response rate was an increasing function of intensity at the high lever position but not at the low position. Correlations between rearing and leverpressing, and between jumping and leverpressing, were not significant for either lever position. Bolles’ (1970) theory of species-specific defense reactions was invoked to account for initial facilitated performance.     

Dominance,the bidirectional hypothesis,and Pavlovian backward conditioning in the US-US paradigm

Based upon considerations raised by Soviet research, the role of relative stimulus intensity, or dominance, in the unconditioned stimulus-unconditioned stimulus (US-US) paradigm was investigated under circumstances presumed favorable to the backward conditioned response (CR). Using the classically conditioned forelimb response of the cat, a brief shock (USD delivered to one forepaw preceded a shock (US2) to the opposite forepaw in paired conditioning fashion; subjects in the control group received explicitly unpaired presentations of the stimuli. Conditioning in both the forward and backward directions was evaluated by the appearance of contralateral CRs on test trials to each of the USs. In Experiment 1, a ratio of the intensities between US1 and US2 of 100:80 was used to create a relative dominance in favor of the backward CR. In addition, to evaluate the suggestion that the appearance of the backward CR is retarded in the Pavlovian paradigm, overtraining was provided to a forward conditioning criterion of 200%. In Experiment 2, the cats were exposed to successive reductions in the intensity of US2 to verify manipulations of dominance reportedly involved in the reactivation of a latent backward CR. Although forward conditioning was readily established to USl, there was no evidence of back-ward conditioning to US2 under any of the conditions.     

Postdiscrimination generalization as a function of interresponse time

Following 20 sessions of variable-interval 20-sec reinforcement in the presence of a single 45-deg line-tilt stimulus, three pigeons were trained to discriminate between line tilts of 45 deg correlated with variable-interval 20-sec reinforcement and line tilts of 15 deg correlated with extinction. A generalization test along the line-tilt dimension was administered following a criterion discrimination performance. Gradients derived in terms of relative frequency of response as a function of line tilt indicated strong external stimulus control and exhibited clear peak shift. From the interresponse time (IRT) distributions generated for responding to each test stimulus, probability of response conditional upon IRT (IRTs/Op) was derived as a joint function of line tilt and IRT. The IRTs/Op functions for responses following IRTs in 0.2-sec-wide classes from 0.2 to 1.0 sec and for responses following IRTs in the interval of 1.0 to 2.0 sec were similar to the relative generalization gradients and also exhibited peak shift. Few IRTs were greater than 2.0 sec. External stimulus control was established over responses terminating IRTs both longer and shorter than 1.0 sec.     

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

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

CS and US duration effects in one-trial simultaneous fear conditioning as assessed by conditioned suppression of licking in rats

In three experiments, rats received a single presentation of an auditory conditioned stimulus (CS) beginning simultaneously with an electric grid-shock unconditioned stimulus (US). Later, the CS was presented while the rats licked a drinking tube for water, and CS-elicited suppression of licking was taken as an index of the excitation conditioned to the CS. It was found that conditioning increased as a joint function of the duration of CS-US overlap and US duration. The evidence suggested that weak conditioning due to a brief CS-US overlap could be increased by extending the US beyond CS termination. Extending CS duration beyond US termination, however, did not strengthen conditioning; indeed, extending the CS 60 sec beyond US termination weakened conditioning significantly. It is suggested that these results shed light on a discrepancy in the recent literature on simultaneous conditioning.     

Stimulus generalization of an illness-induced aversion to different intensities of colored water in Japanese quail

After quail (Coturnix coturnix japonica) experienced a toxicosis following ingestion of water colored dark blue or light blue, the stimulus generalization of the aversion to light, medium, and dark blue water was investigated. Birds made ill following ingestion of dark blue water demonstrated a reduced aversion when tested with intensities of blue water weaker than the conditioning intensity. Birds made ill following ingestion of light blue water showed stronger aversions as a function of increasing test intensity, suggesting a stimulus intensity dynamism effect. The stimulus generalization and stimulus-intensity dynamism effects observed in this study where visual cues were used is in agreement with other results within the illness-induced aversions paradigm where rats were tested with varying concentrations of a gustatory stimulus.     

Conditioned suppression of a positively reinforced shuttle response

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

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.     

Solving Pavlov's puzzle: Attentional, associative, and flexible configural mechanisms in classical conditioning

This article introduces a new "real-time" model of classical conditioning that combines attentional, associative, and "flexible" configural mechanisms. In the model, attention to both conditioned (CS) and configural (CN) stimuli are modulated by the novelty detected in the environment. Novelty increases with the unpredicted presence or absence of any CS, unconditioned stimulus (US), or context. Attention regulates the magnitude of the associations CSs and CNs form with other CSs and the US. We incorporate a flexible configural mechanism in which attention to the CN stimuli increases only after the model has unsuccessfully attempted learn input-output combinations with CS-US associations. That is, CSs become associated with the US and other CSs on fewer trials than they do CNs. Because the CSs activate the CNs through unmodifiable connections, a CS can become directly and indirectly (through the CN) associated with the US or other CSs. In order to simulate timing processes, we simply assume that a CS is formed by a temporal spectrum of short-duration CSs that are activated by the nominal CS trace. The model accurately describes 94?% of the basic properties of classical conditioning, using fixed model parameters and simulation values in all simulations.     

Effects of proximal unconditioned stimulus preexposure on ingestional aversions learned as a result of taste presentation following drug treatment

Taste aversions were conditioned by exposing subjects to a 1.0% saccharin solution 30 min after an injection of lithium chloride. The aversion learning was disrupted if subjects had also received an additional lithium injection some time earlier (Experiments 1–3). This interference effect of US preexposure was a decreasing function of the preexposure interval, beyond the optimal interval (105 min) for observing the phenomenon (Experiment 1), and was directly related to the dose of the preexposure injection (Experiment 2). No interference with conditioning occurred at short (e.g., 30-min) preexposure intervals (Experiment 1), probably because under these circumstances the preexposure injection itself conditioned a strong aversion (Experiment 4). At moderate (105-min) but not at short (30-min) preexposure intervals, the interference with aversions learned as a result of taste exposure following drug injection was comparable to the interference with learning in a more conventional forward conditioning procedure (Experiments 3 and 4). These findings are similar to previously documented effects of proximal CS- and US-preexposure and are consistent with recent stimulus rehearsal and opponent-process theories.     

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.     

Characteristics of the lithium-mediated proximal US-preexposure effect in flavor-aversion conditioning

Taste-aversion learning in rats is disrupted if the subjects are exposed to the unconditioned stimulus (US) shortly before the conditioning trial but not if this single US preexposure treatment occurs 1 day or more before conditioning. Several characteristics of this proximal US-preexposure phenomenon were explored. Experiment 1 showed that the time course of the interference with conditioning is directly related to the preexposure drug dose. Experiment 2 demonstrated that the interference effect is evident even if the test for aversion learning is conducted following a drug injection, thereby minimizing stimulus generalization decrement for the preexposed subjects. Finally, Experiment 3 showed that disruption of the contingent relationship between tastes and drug effects is probably not responsible for the proximal US-preexposure phenomenon because the interference with conditioning occurs regardless of whether or not the preexposure drug treatment is paired with a novel flavor. These findings, together with previous research, demonstrate the remarkably robust character of the proximal US-preexposure phenomenon.     

Conditioned suppression is an inverted-U function of footshock intensity

Most theories of associative learning assert that conditioned responding to a target cue is a monotonically increasing function of unconditioned-stimulus (US) intensity. In a lick suppression preparation with rats, a cue was paired with a 0.4-, 0.6-, 0.8-, 1.0-, 1.2-, or 1.4-mA footshock in Experiment 1a, and with a 0.3-, 0.8-, 1.3-, or 1.8-mA footshock in Experiment 1b. Subsequent suppression in response to the cue was an inverted-U function of the US intensity. In Experiment 2, we demonstrated that massive extinction of the training context and compound conditioning can each attenuate the response decrement caused by training with a high-intensity US. The sometimes-competing-retrieval model (Stout & Miller, Psychological Review 114:759–783, 2007) provides a better fit to these data than do several other models of associative learning.     

On the generality of conditioned bradycardia in rabbits: Assessment of CS and US modality

To determine whether the magnitude of heart rate (HR) slowing induced by classical conditioning contingencies is comparable under a broad range of stimulus conditions, experiments were conducted in which rabbits were exposed to tones, increases in illumination, or vibratory stimuli as conditioned stimuli (CSs) and in which paraorbital electric shocks, corneal airpuffs, or intraoral pulses of water served as unconditioned stimuli (USs). The results indicated that conditioned bradycardia was elicited by all three CSs. Moreover, when a corneal airpuff served as the US, small but reliable CS-evoked HR decelerations also occurred. Finally, CS-evoked HR slowing also occurred in response to a tone CS employed in an appetitive task, in which water was the US. These findings suggest that HR slowing is a general phenomenon that occurs when rabbits are exposed to signals that systematically predict aversive or appetitive consequences according to a Pavlovian conditioning paradigm.     

Effects of expected vs. unexpected proximal US preexposure on taste-aversion learning

Animals were first conditioned to expect lithium treatment following exposure to one taste solution (the CS+) and to expect no drug treatment following exposure to another flavor (the CS?). All subjects then received a saccharin taste-aversion conditioning trial. In Experiment 1, this conditioning trial was preceded 0, 1, 2, 4, or 6 h earlier by exposure to the CS+ flavor for independent groups. The CS+ exposure attenuated saccharin aversion learning if it occurred immediately before the saccharin conditioning trial but not if it occurred 1 h or more before conditioning. In Experiment 2, the saccharin conditioning trial was preceded 3 or 4.5 h earlier by a lithium injection. This proximal US preexposure injection was either unannounced (Li) or preceded by exposure to the CS+ (CS+Li) or the CS? (CS?Li) stimuli. The US preexposure attenuated saccharin aversion learning in all cases. However, the interference effect was less when the preexposure injection was expected (CS+Li) than when it was unexpected (CS?Li). This outcome could not be explained in terms of direct effects of the CS+ and CS? stimuli on the saccharin conditioning trial, and shows that the proximal US preexposure effect is a function of not only the drug dosage and preexposure interval, but also the anticipation of the drug pretreatment.     

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.