Fear-potentiated startle using three conditioned stimulus modalities

The capacities of three different conditioned stimulus modalities (light, noise, and airflow produced by a fan) to produce fear-potentiated startle were evaluated. Previous experiments have shown that following either light-shock or noise-shock pairings, both the light and noise conditioned stimuli acquire the ability to potentiate the acoustically elicited startle response in rats (the so-calledfear-potentiated startle effect). In Experiment 1, the ability of airflow produced by a fan to act as a conditioned stimulus was investigated. Rats were given either paired or impaired fan-shock training followed by a test for fear-potentiated startle. The fan conditioned stimulus potentiated startle only in the group given explicit fan-shock pairings. In Experiment 2, we evaluated the discriminability of the three conditioned stimulus modalities. Rats were given light, noise, or fan-shock pairings and were subsequently tested for fear-potentiated startle with the trained conditioned stimulus as well as the two remaining novel conditioned stimuli. Only the trained conditioned stimulus potentiated startle. These results show that fear-potentiated startle can be produced with three discriminable conditioned stimulus modalities, allowing the future use of fear-potentiated startle in the investigation of higher order conditioning phenomena.     

Extinction and food-reinforced inhibition of conditioned salivation in dogs

The present experiment compared two methods of eliminating a classically conditioned response in dogs, extinction and reinforcement of nonsalivation, using both a within- and between-subjects experimental design. Eighteen dogs were trained for 16 days in Phase I, 16 days in Phase II, and 8 days in Phase III. In Phase I, each subject received classical conditioning training to two stimuli. In Phase II, Group 1 received extinction training to one stimulus and reinforcement of nonsalivation to the other stimulus. Group 2 received continued classical conditioning training to one stimulus and reinforcement on nonsalivation training to the other. Group 3 received continued classical conditioning training to one stimulus and extinction training to the other. In both the within- and between-subjects comparisons, responding to the stimulus associated with extinction was eliminated faster than responding to the stimulus associated with reinforcement of nonsalivation.     

Conditioned inhibition of analgesia

Stimuli that predict the occurrence of aversive events come to elicit conditioned analgesia. Experiments 1A and 1B examined the possibility that conditioning can inhibit analgesia when stimuli are paired in a backward fashion with a shock US (Pavlovian CS- s). Analgesia conditioned in response to shock context exposure was reversed during the CS- (light) presentation after four sessions. The ability of the CS- to function as a conditioned inhibitor of analgesia was then evaluated in both summation (Experiment 1A) and retardation-of-acquisition testing (Experiments 1A and 1B). The results support the conclusion that a stimulus presented after shock in a backward fashion comes to be a conditioned inhibitor of analgesia. Experiments 2A and 2B examined the assumption that the results obtained with our pain sensitivity measure (tailflicking in response to radiant heat) reflect changes in responsiveness to painful input, rather than a general motor inhibition or general insensitivity to sensory input. In Experiment 2A, tailflick responding to painful and nonpainful input was compared in animals receiving either morphine or saline. In Experiment 2B, tailflick responding to painful and nonpainful input to the tail was compared in both the shock and a neutral context. In both experiments, only the painful input yielded changes in responsivity. The results support the conclusion that the alterations in pain sensitivity produced by the CS- for shock represents a conditioned inhibition specific to pain.     

Stimulus duration and conditioned reinforcing value measured by a learning-tests procedure

The relationship between the duration of stimuli and their conditioned reinforcing effect was investigated using a learning-tests procedure. In Experiment 1, stimuli were the same duration on training (stimulus → reward) and test (choice response → stimulus). Ten- and 30-sec stimuli provided effective differential conditioned reinforcement but 3-sec stimuli did not. In Experiment 2, different pigeons had each combination of the 3- and 30-sec stimuli on training and test trials. Evidence of conditioned reinforcement was obtained only for the birds with 30-sec stimuli on both training and test. The results were interpreted as indicating that stimuli become effective conditioned reinforcers on test trials only when their duration exceeds the duration of differential short-term memory cues resulting from a difference in the events that precede them on training and test trials.     

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.     

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.     

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.     

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.     

Behavioral and associative effects of differential outcomes in discrimination learning

The role of the reinforcer in instrumental discriminations has often been viewed as that of facilitating associative learning between a reinforced response and the discriminative stimulus that occasions it. The differential-outcome paradigm introduced by Trapold (1970), however, has provided compelling evidence that reinforcers are also part of what is learned in discrimination tasks. Specifically, when the availability of different reinforcing outcomes is signaled by different discriminative stimuli, the conditioned anticipation of those outcomes can provide another source of stimulus control over responding. This article reviews how such control develops and how it can be revealed, its impact on behavior, and different possible mechanisms that could mediate the behavioral effects. The main conclusion is that differential-outcome effects are almost entirely explicable in terms of the cue properties of outcome expectancies—namely, that conditioned expectancies acquire discriminative control just like any other discriminative or conditional stimulus in instrumental learning.     

Protection from extinction provided by a conditioned inhibitor

Three conditioned suppression experiments with rats as subjects investigated the influence of higher order associations in determining the response potential of a target stimulus. In these experiments, a Pavlovian conditioned inhibitor was compounded with the target cue during extinction treatment. In Experiment 1, strong suppression was observed to the target cue that was given extinction treatment in the presence of a conditioned inhibitor, relative to a target that was extinguished with an associatively neutral cue or was extinguished alone, suggestive of enhanced protection from extinction provided by a conditioned inhibitor. This effect was replicated in a sensory preconditioning preparation in Experiment 2; in Experiment 3, in a sensory preconditioning preparation, this protection effect was retroactively attenuated when the conditioned excitor used to train the conditioned inhibitor was extinguished following extinction of the target. This provides evidence that, at least in a sensory preconditioning preparation, stimuli that are only indirectly associated with the target cue can contribute to the response potential of that target.     

Effects of stimulus spacing on steady state gradients of inhibitory stimulus control

When the response of pigeons is maintained to a number of stimulus wavelengths, but extinguished to one (S?), the birds peck more rapidly at stimuli near the S? than at more distant stimuli. The present study explores this “dimensional contrast” effect as a function of the number and spacing of test wavelengths. A fixed portion of the wavelength continuum was spanned by 5, 9, 13, or 49 stimuli, which appeared in random sequence behind a standard pecking key. At the end of each 20-sec trial, pecks to test stimuli produced a conditioned reinforcer (sometimes followed by food), while pecks to the S? stimulus produced only darkness. Dimensional contrast “shoulders” developed to test stimuli on either side of the S?; these shoulders were of approximately the same height and wavelength position for all but the 5-stimulus (widely spaced) condition, and were comparable to the original contrast results with 25 stimuli. The results strongly suggest that the extent and locus of contrast shoulders are largely independent of the number and spacing of test stimuli.     

Evaluating conditioning of related and unrelated stimuli using a compound test

Three experiments used a compound test procedure to evaluate whether superior conditioning results from the pairing of stimuli that are related to each other. In each case, a stimulus compound was tested after its component conditioned stimuli (CSs) had been conditioned by the same unconditioned stimuli (USs) arranged such that either related or unrelated CSs and USs were paired. Experiment 1 explored auditory and gustatory stimuli conditioned by LiCl or shock, using rats. Experiments 2 and 3 used second-order conditioning in pigeons to pair stimuli that were similar by virtue either of qualitative features or of shared physical location. In each case, the compound test provided clear evidence that pairing related stimuli produces superior associative learning.     

Potentiation of responding on a VR schedule by a stimulus correlated with reinforcement: Effects of diffuse and localized signals

In Experiment 1, rats were trained to leverpress on a variable ratio (VR) 30 schedule with a 500-msec delay between the reinforced response and food delivery. Subjects that experienced a signal during the delay responded faster than did control subjects that received the stimulus un-correlated with reinforcement. Higher response rates were obtained when the stimulus used to signal reinforcement was auditory rather than visual. Experiments 2 and 3 compared the effects of signaling reinforcement with either a localized or a diffuse light on responding maintained by VR schedules of reinforcement. Elevated response rates were observed with the diffuse stimulus, but the localized stimulus failed to produce such potentiation. Experiment 3 also examined the conditioned reinforcing power of localized and diffuse visual stimuli. These results are discussed with reference to (1) theories of selective association and sign tracking and (2) their implications for current theories of signaling reinforcement.     

Blocking of the sexual conditioning of differentially effective conditioned stimulus objects

The blocking phenomenon was investigated in the sexual response system of male Japanese quail. Access to a live female quail served as the unconditioned stimulus (US). The same audiovisual cue served as the pretrained stimulus in all of the experiments. Following asymptotic conditioning of the audiovisual cue, a second conditioned stimulus (CS2) was added. In Experiment 1, CS2 was a rectangular wood block that had little or no resemblance to a female quail and could not support copulatory behavior. In Experiment 2, CS2 was a terrycloth object that had no quail parts but could support copulatory behavior, and, in Experiment 3, CS2 was a terrycloth object that had a taxidermically prepared head of a female quail added. The terrycloth-only object supported more rapid conditioning than did the wood block, but the blocking effect was obtained with both kinds of stimuli. Approach responding to the terrycloth + head object required pairing it with copulatory opportunity, and the terrycloth + head object supported at least as rapid conditioning as did the terrycloth-only object. However, responding to the terrycloth + head object was not blocked by the pretrained audiovisual cue. These results indicate that the blocking effect occurs in sexual conditioning even with stimulus objects that can support copulation. However, the addition of species-typical head cues to an object makes that object such a powerful stimulus that conditioned approach responding to it cannot be blocked by a previously conditioned arbitrary audiovisual cue.     

Microanalysis of variable-interval performance during stimulus compounding

Additive summation is observed when more responses are emitted to the simultaneous presentation (tone-plus-light) of independently conditioned stimuli (tone and light) than to either stimulus presented alone. The current experiment sought to determine if this increased rate during tone-plus-light was a function of a new modal interresponse time (IRT) or a differential mixing of pauses with a modal IRT characteristic of the responding in tone and light alone. Three rats were trained on a three-component multiple schedule where tone and light were each associated with a variable-interval 30-sec schedule while a variable-interval 100-sec operated in the simultaneous absence of these stimuli, tone-off and light-out. Baseline response rates were 2–4 times as high in tone or light as in their absence. In testing, more responses were emitted to tone-plus-light than to tone or light by all animals, but the modal IRT was in the 0.2–0.4-sec IRT bin for all test conditions. Tone-plus-light controlled fewer long IRT values and more responses in the short modal category than tone or light alone. These results support the response mixing hypothesis of stimulus control; i.e., no “new” behavior was observed during the novel combination of stimulus elements, only a mixture of previously reinforced behavior patterns in different proportions.     

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.     

Summation of undetected excitation following extinction of the CER

The effects of compounding two conditioned stimuli (CSs), each of which had been extinguished to varying degrees in different groups of rats given identical acquisition training, was examined within the conditioned emotional response paradigm. Greater suppression to the compound than to the individual CSs was observed following 6, 12, 48, 72, and 96 stimulus extinction trials, although after only 12 extinction trials suppression to the individual stimuli was no longer observed. The amount of compound suppression decreased progressively as the number of extinction trials increased until, after 120 extinction trials on each stimulus, the compound no longer elicited observable suppression. Control group data indicated that the observed summation effect could not be attributed to disinhibition. The possible role of the summation of undetected excitation in studies examining configurai conditioning, avoidance conditioning and reinstatement is discussed briefly.     

Chemosensory-based contextual conditioning inHermissenda crassicornis

In two experiments, the marine molluskHermissenda crassicornis was exposed to context discrimination training. In one context, defined by the presence of a diffuse chemosensory stimulus (shellfish extract A), brief, unsignaled, unconditioned stimuli (USs; high-speed rotation) were presented; in a second context, defined by the presence of shellfish extract B, no USs were presented. Animals were then tested (at both 1.5 and 24 h) by exposing them to small pieces of the shellfish meat used to define the two contexts. The latency to strike at the meat served as an index of the context-US association. In Experiment 1, the latency to strike at the cue associated with rotation was reduced relative to both preconditioning strike latencies and the associatively neutral cue. However, in a two-choice test where the animals could approach the conditioned or neutral stimulus, the animals regularly avoided the stimulus paired with rotation. Moreover, if, following conditioning, the animals were presented with an unsignaled rotation in the conditioned context or the neutral context, the animals exhibited more effective defensive clinging (an unconditioned reflex normally elicited by rotation) in the conditioned context, suggesting that it “prepared” the animal for the aversive US. In total, these results demonstrate thatHermissenda is capable of making associations to diffuse background (contextual) stimuli. Moreover, the results suggest that pairing the chemosensory cue with an aversive US elicits a strike response inHermissenda when the animal is placed in forced contact with the cue and an active avoidance response when the animal can choose between that cue and a neutral cue.     

A behavior systems view of responding to probe stimuli during an interfood clock

Two experiments used a behavior systems approach to relate the form of responses during an interfood clock to the temporal distance of the individual clock stimuli to food. Stimuli proximate to food should better control a focal search mode and related responses, whereas stimuli temporally distant from food should better control a general search mode and related responses. Experiment 1 conditioned two groups of rats with a sequence of four equal-length 12-sec clock stimuli that terminated with food and then tested for the conditioning of a general search mode by presenting an unconditioned moving probe stimulus (either a rolling ball bearing or a rotating mechanical door) during each of the clock stimuli. Consistent with a behavior systems view, contact with the ball bearing was markedly greater during a clock stimulus distant from food. The absence of similar differential contact of the door across the clock stimuli showed that the effect was specific to the ball bearing rather than a general response to stimulus dimensions of movement and sound. Experiment 2 showed that the general search mode was controlled by the clock stimulus rather than the passage of time.     

On the nature of CS and US representations in Pavlovian learning

A significant problem in the study of Pavlovian conditioning is characterizing the nature of the representations of events that enter into learning. This issue has been explored extensively with regard to the question of what features of the unconditioned stimulus enter into learning, but considerably less work has been directed to the question of characterizing the nature of the conditioned stimulus. This article introduces a multilayered connectionist network approach to understanding how “perceptual” or “conceptual” representations of the conditioned stimulus might emerge from conditioning and participate in various learning phenomena. The model is applied to acquired equivalence/distinctiveness of cue effects, as well as a variety of conditional discrimination learning tasks (patterning, biconditional, ambiguous occasion setting, feature discriminations). In addition, studies that have examined what aspects of the unconditioned stimulus enter into learning are also reviewed. Ultimately, it is concluded that adopting a multilayered connectionist network perspective of Pavlovian learning provides us with a richer way in which to view basic learning processes, but a number of key theoretical problems remain to be solved, particularly as they relate to the integration of what we know about the nature of the representations of conditioned and unconditioned stimuli.