The effects of d-amphetamine on the temporal control of operant responding in rats during a preshock stimulus.

The operant behavior of six rats was maintained by a random-interval schedule of reinforcement. Three-minute periods of noise were superimposed on this behavior, each period ending with the delivery of an unavoidable shock. Overall rates of responding were generally lower during the periods of noise than in its absence (conditioned suppression). These suppressed response rates also exhibited temporal patterning, with responding becoming less frequent as each noise period progressed. The effects of d-amphetamine on this behavioral baseline were then assessed. In four animals the relative response rates during the noise and in its absence suggested that the drug produced a dose-related decrease in the amount of conditioned suppression. However, this effect was often due to a decrease in the rates of responding in the absence of the preshock stimulus, rather than to an increase in response rates during the stimulus. Temporal patterning in response rates during the preshock stimulus was abolished, an effect that was interpreted in terms of rate-dependent effect of d-amphetamine. This study thus extends rate-dependent analyses of the effects of amphetamines to the patterns of operant behavior that occur during a preshock stimulus, and which have been discussed in terms of the disrupting effects of anxiety on operant behavior.     

Response rate, reinforcement frequency, and conditioned suppression

In the first of two experiments, periods of noise were terminated with unavoidable shock to 36 rats. The rats' continuously reinforced responding was later completely suppressed during the noise when it was introduced without shock. The rats were then assigned to nine experimental groups. Each group was exposed to different paced variable-interval schedules of reinforcement, which independently controlled response rate and reinforcement frequency. Periods of the noise were periodically superimposed on these schedules, and loss of response suppression was studied. Differences between the groups were assessed statistically. The second experiment used a steady-state design. Six rats were exposed to paced schedules which generated two alternating response rates but gave constant reinforcement frequencies, and six rats to schedules which maintained the same response rates throughout, but in which the reinforcement frequency was alternately high and low. Response suppression was studied during a pre-shock stimulus superimposed on each rat's two behavioral baselines. Both experiments suggest that (1) conditioned suppression is affected by rate of operant responding, high rates being most suppressed, and (2) the frequency of reinforcements obtained also has an effect, most suppression occurring when frequency is low.     

Conditioned suppression of an avoidance response by a stimulus paired with food

Three food-deprived Long-Evans rats were exposed to a non-discriminated shock avoidance procedure. Superimposed upon this operant avoidance baseline were periodic presentations of a conditioned stimulus that was paired with food, the unconditioned stimulus. These pairings resulted in increases in the rate of shock over that recorded when the conditioned stimulus was not present. A traditional suppression ratio failed to reveal any differential effect of the conditioned stimulus on the overall rate of avoidance responding, although all subjects showed a consistent pattern of pausing and postshock response bursts during presentations of the conditioned stimulus. When food was withheld during a final extinction phase, the conditioned stimulus ceased to occasion increases in shock rates and disruptive postshock response bursts were eliminated. An analysis of conditioned suppression procedures is proposed that stresses not only operant-Pavlovian or appetitive-aversive incompatibility, but also the manner in which the baseline schedule of reinforcement affects operant behavior changes that are elicited by the superimposed Pavlovian procedure.     

Conditioned suppression and conditioned enhancement with the same positive UCS: an effect of CS duration

Previous experiments have shown that positively reinforced operant responding is suppressed during a conditioned stimulus terminated with an electric shock (conditioned suppression). In the present experiment, the conditioned stimulus was terminated with a positive unconditioned stimulus, and it was found that the duration of the conditioned stimulus was a key factor in determining whether response suppression or response enhancement was observed during the stimulus. The lever-pressing responses of rats were maintained by a variable-interval schedule of food reinforcement. While the rats were pressing the lever, a light was occasionally turned on, its offset coincident with a brief period of access to a sucrose solution. In consecutive blocks of sessions, the light duration was 40 sec, 12 sec, or 120 sec. Results showed that the rate of lever pressing was substantially suppressed during the 12-sec stimulus, slightly suppressed during the 40-sec stimulus, and enhanced during the 120-sec stimulus.     

Some effects of the conditioned suppression paradigm on operant discrimination performance

Three experiments were conducted with rats to determine the effects of electric shock on responding during an operant discrimination. In two of these experiments, a conditioned suppression procedure was superimposed upon a stimulus signalling the availability of food reinforcement (S(D)). Response rates were greatly suppressed, not only in the warning signal periods which preceded each shock, but in the presence of S(D), and the stimulus signalling the unavailability of reinforcement (S(Delta)) as well. A third experiment, in which a very mild shock was used without a warning signal, demonstrated an increased response rate in S(D) and S(Delta), although this effect was rather unsystematic. In a similar study, Hearst (1965) found an increased rate in S(Delta) independent of any change in the S(D) rate. The present study failed to obtain Hearst's effect but illustrated a suppressive effect with a similar procedure.     

Positive conditioned suppression: effects of CS duration

During a brief conditioned stimulus (15 or 30 sec) that terminated with the response-independent delivery of banana pellets, operant responding reinforced by other food pellets according to a variable-interval schedule of reinforcement was suppressed in the squirrel monkey. Conditioned stimuli of longer duration (1, 2, and 3 min) did not reliably affect the rate of operant performance. Brief conditioned stimuli generated homogeneous response patterns of nearly complete suppression. Increasing the CS duration did not enhance responding, as previously reported, but led to alternate bursting and pausing, which suggested a loss of control by the conditioned stimulus. The results suggest that the magnitude of "positive" or "negative" conditioned suppression reflects the strength of the classical conditioning process.     

A conditioned reinforcer maintained by temporal association with the termination of shock

Two experiments were conducted to determine whether a stimulus can be established as a positive conditioned reinforcer by associating it with the termination of shock, but without training the animal to make any response in its presence. In the first, six rats were conditioned to press a bar to terminate shock on a variable ratio schedule; white noise was then substituted as the immediate consequence, with the shock terminating 30 sec after the last press in its presence. It was found that the rate of pressing in the absence of noise depended on the contingency between the pressing and the noise. The second experiment sought to determine whether the difference in rates before and after the onset of the noise was due to the reinforcement of prior responding by the onset of the noise or to the suppression of subsequent responding by differential reinforcement of competing behavior. Six more rats were trained in the same manner, but with shock terminating 30 sec after the onset of the noise, regardless of what the animal did in its presence. Again the rate was higher before the onset of the noise, indicating that pressing was indeed maintained by the noise as a conditioned reinforcer.     

Food and intracranial stimulation responding suppressed with regular-interval shock.

Attenuation of conditioned suppression during intracranial stimulation was compared with that during food reinforcement. Response rates controlled by food and by brain stimulation were equalized on a multiple schedule by adjusting the stimulating current. When foot shock was delivered during timeout periods separating response components, responding for food was significantly more suppressed than responding for brain stimulation. When components were shortened from 10 to 2 minutes, responding maintained by either food or brain stimulation showed a similar temporal pattern of suppression preceding each shock, but responding in the component involving food remained significantly more suppressed. Explanations for the attenuated suppression during brain stimulation based on neural disruption, stimulus blocking, and analgesic properties were questioned. The increased responding during brain stimulation seemed to reflect greater response strength relative to food reinforced responding.     

Temporal properties of responding during stimuli that precede response-independent food

Pigeons' keypecks were reinforced with grain on the average of once per minute by schedules that maintained low response rates and by schedules that maintained high response rates. During these schedules, a fixed-duration conditioned stimulus (CS) ranging from 7.5 to 120 sec in duration across conditions terminated with response-independent food. Response rates during the CS were inversely related to CS duration. The rates and the temporal patterns of responding during the shortest CS were similar whether the ongoing schedule maintained high response rates or low response rates. As CS duration increased, the rate and pattern of responding during the CS converged on the rate and pattern of responding maintained by the baseline schedule. These data indicate that changes in responding during stimuli that signal response-independent reinforcement are not homogeneous throughout the CS; that response measures, such as “suppression ratios”, which presume homogeneity may mislead us; and that conditioned suppression and conditioned enhancement may be better talked about in terms of species-specific approach and avoidance than in terms of emotional states.     

Conditioned suppression or facilitation as a function of the behavioral baseline

Rats were exposed to a multiple schedule of reinforcement. During one component, a bar-press was followed by reinforcement only if it occurred between 15 and 20 sec after the previous response. This differential-reinforcement-of-low-rate (DRL) schedule produced a typical slow rate of responding. During the other component, reinforcement followed the first response to be emitted during limited periods of time which occurred at fixed intervals. These fixed-interval schedules with a limited hold produced higher response rates, described as `interval' or `ratio-like' behavior. Responding during the DRL component increased in frequency during a tone which ended with an unavoidable shock of low intensity, but decreased during the tone when the shock intensity was raised. The `interval' and `ratio-like' responding decreased in frequency during the tone at all shock intensities. Initial acceleration of the DRL responding appeared to be due to adventitious punishment of collateral behavior which was observed between the bar-presses. The more severe conditioned suppression during the fixed-interval components might be the result of the lower probability of reinforcement after any single response.     

Summation of excitatory and inhibitory control produced by traditional tone-shock contingencies and autocontingencies

Previous research has demonstrated that rats can use unsignaled shock to predict subsequent periods free from shock. This shock-no shock stimulus arrangement, termed an autocontingency, has appeared less likely to exert behavioral control when a traditional tone-shock contingency was simultaneously available. The present research examined the generality of CS-US contingency dominance in a conditioned suppression paradigm by using a summation test in which “probe” stimuli derived from tone-shock contingencies were superimposed upon responding maintained by an autocontingency. In experiment 1, an inhibitory CS accelerated responding only when responding was normally suppressed by the autocontingency. In experiment 2, an excitatory CS failed to yield conditioned suppression during an inhibitory (accelerative) period produced by the autocontingency. Unlike our previous findings (e.g., Davis, Memmott & Hurwitz, 1975), these results do not support a general notion of tone-shock contingency dominance over autocontingencies. Behavioral control by autocontingencies appears robust and “holds its own” in summation with both excitatory and inhibitory CSs derived from traditional contingencies.     

Punishment as a discriminative stimulus and conditioned reinforcer with humans

Mental hospital patients were reinforced for responding in a two-response operant situation. When a noise was used to punish one of the responses, all subjects shifted to the unpunished one. When the noise was then paired with positive reinforcement, the subjects responded to produce the noise. Also, a novel response was reinforced by noise in the absence of other reinforcers. This study with humans extends the findings of previous studies with animals in revealing how a punishing stimulus can acquire discriminative or conditioned reinforcing properties.     

Enhancement of conditioned autonomic responses in monkey when preshock signals occasion operant suppression.

Classical pairings of a sound stilulus with shock elicited larger magnitude and more rapidly conditioned autonomic responses when subjects were responding on variable-interval schedules for food than when they were eating freely available food. The difference was not attributable to changes in control values of heart rate and blood pressure, or to alterations in motor activity, but appeared related to operant suppression.     

Sexual reinforcement in the female rat.

Sexual reinforcement in the female rat was studied in a preparation that allowed continuous operant responding for access to a male rat leading to intromission. Experiment 1 used a high operant level nose-poke response to test the possible reinforcing effects of some components of access to a male. A simple tone stimulus used as a conditioned reinforcer and two odor stimuli, target male bedding and emulsified preputial gland, were tested. None of these contingent events altered responding above or below operant level. Access to the male, which was always accompanied by intromission, immediately increased response rate when it was made contingent upon the nose-poke response. Performance on fixed-ratio schedules was erratic, and response rate was low in comparison to typical food-reinforced responding. An interresponse-time analysis indicated, however, that some effect of the ratio contingency may have been present. In Experiment 2, several modifications of the procedure were tested with the objective of creating a more tractable preparation for behavior analysis. Response type and the hormone delivery method were changed, and 2 target males were used instead of 1. The latter tripled the average number of reinforcers earned in a single session. Differences between sexual and other reinforcers are discussed in terms of procedural, quantitative, and motivational aspects of the sexual reinforcement procedure.     

Conditioned suppression of an avoidance response

A signal followed by shock was presented at irregular intervals during a free-operant avoidance schedule. The effects of this procedure were studied in terms of the rate of unavoided shock in the presence and absence of the signal and the rate of response before and during the signal. Three shock intensities were employed. Response enhancement as well as response suppression were observed; irrespective of changes in responding, shock rates substantially increased during signalled periods compared to non-signalled periods. Shock rates in non-signalled periods were generally higher than during training.     

Effects of signals preceding and following shock on baseline responding during a conditioned-suppression procedure

Long-Evans rats were exposed to a succession of conditioned-suppression procedures involving pairings of (1) signal-shock, (2) shock-signal, and (3) a signal-shock-signal sequence in which first and second signals were at first physically identical. Traditional suppression of food-reinforced responding was obtained under the signal-shock arrangement, and exposure to the shock-signal sequence resulted in conditioned enhancement of responding during the signal. The signal-shock-signal condition reliably suppressed responding during the first signal, but produced no differential effect on response rate during the second signal. Baseline responding was least changed from preshock rates under the signal-shock-signal procedure, but baseline rate was considerably reduced under the signal-shock and shock-signal arrangements, the latter yielding most substantial reductions. A second experiment indicated that the magnitude and direction of changes in baseline responding reported in Experiment I were not confined to cases in which the first and second signals in the signal-shock-signal arrangement were physically identical. It is suggested that the major effects of the conditioned-suppression procedure on response rate might not be confined to presentations of the signal.     

Blocking and nonsimultaneous compounds: Comparison of responding during compound conditioning and testing

Three blocking experiments were run using a conditional emotional response procedure with rats as subjects. In each experiment, following initial conditioning to a light stimulus, blocking rats were conditioned to a compound in which a short-duration (30-second) noise was superimposed on the terminal portion of the longer-duration light. In Experiment 1, the light was always three minutes in duration; in Experiment 2, the light was switched from a constant (three-minute) to a variable (0.5–5.0 minute) duration at the start of compound conditioning; in Experiment 3, the light was variable in duration throughout conditioning. Control rats received the same compound conditioning experiences as the blocking rats but did not receive prior conditioning to the light by itself. These temporal manipulations had strong and systematic influence on the rats’ pattern of responding during compound conditioning. Most notably, experience with a constantduration light resulted in both blocking and compound animals showing little conditional suppression to the early conditional stimulus portion of light by itself, followed by strong conditional suppression to the terminal 30 seconds of light plus noise. When testing was done with the noise in isolation subsequent to compound conditioning, however, an equally strong blocking effect was obtained across all the temporal manipulations. In all cases, the blocking animals showed much less conditional suppression to the noise than did the compound control animals. This lack of correspondence between the rats’ responding during compound conditioning and their response to the noise by itself is theoretically puzzling. The confounding effect of inhibition of delay may provide at least a partial explanation of this pattern of results.     

Operant and nonoperant vocal responding in the mynah: Complex schedule control and deprivation-induced responding

Several recent studies have been concerned with operant responses that are also affected by nonoperant factors, (e.g., biological constraints, innate behavior patterns, respondent processes). The major reason for studying mynah vocal responding concerned the special relation of avian vocalizations to nonoperant emotional and reflexive systems. The research strategy was to evaluate operant and nonoperant control by comparing the schedule control obtained with the vocal response to that characteristic of the motor responses of other animals. We selected single, multiple, and chain schedules that ordinarily produce disparate response rates at predictable times. In multiple schedules with one component where vocal responding (“Awk”) was reinforced with food (fixed-ratio or fixed-interval schedule) and one where the absence of vocal responding was reinforced (differential reinforcement of other behavior), response rates never exceeded 15 responses per minute, but clear schedule differences developed in response rate and pause time. Nonoperant vocal responding was evident when responding endured across 50 extinction sessions at 25% to 40% of the rate during reinforcement. The “enduring extinction responding” was largely deprivation induced, because the operant-level of naive mynahs under food deprivation was comparable in magnitude, but without deprivation the operant level was much lower. Food deprivation can induce vocal responding, but the relatively precise schedule control indicated that operant contingencies predominate when they are introduced.     

Increased reinforcement when timeout from avoidance includes access to a safe place

Three experiments investigated the reinforcing value of access to a safe place during timeout from an avoidance schedule. Rats were trained on conjoint schedules in which responding both postponed shock on a free-operant avoidance schedule and produced periods of timeout on fixed-ratio schedules. In some conditions, a shelf was inserted into the operant chamber during timeout, enabling subjects to get off the grid floor. The combination of timeout and shelf maintained substantially higher response rates than the baseline avoidance schedule with ratio requirements as high as 90 (Experiment I). Adding the shelf to timeouts in one component of multiple fixed-ratio schedules of timeout resulted in higher response rates in the component where the shelf was included (Experiment II). When timeouts with and without the shelf were arranged on concurrent schedules, the shelf-timeout combination was preferred, even when of shorter duration than timeout alone (Experiment III). In all three experiments, subjects climbed on the shelf, although all shocks were cancelled during timeout periods. The results could not be accounted for solely in terms of the reinforcing properties of changes in shock rates, but required an interpretation that ascribed conditioned reinforcing value to stimuli associated with such changes.     

CHANGES IN BLOOD PRESSURE AND HEART RATE DURING FIXED‐INTERVAL RESPONDING IN SQUIRREL MONKEYS

Episodic and sustained increases in heart rate and mean arterial blood pressure can occur with recurring patterns of schedule‐controlled behavior. Most previous studies were conducted under fixed‐ratio schedules, which maintained a consistent high rate of responding that alternated with periods of no responding during times when the schedule was not in operation. The present study examined changes in heart rate and blood pressure under fixed‐interval schedules which maintained a range of rates that varied from little or no responding at the beginning of the fixed interval to high rates at the end of the interval. The relations of cardiovascular function to rate of responding were examined. Squirrel monkeys prepared with arterial catheters were trained to respond under fixed‐interval schedules of electric‐shock presentation. The duration of the interval was varied across sessions and cardiovascular parameters were examined. Local rates of responding were typically near zero during timeout periods, low at the beginning of each fixed‐interval cycle, and then increased as the fixed interval progressed. At most schedule durations, arterial blood pressure and heart rate levels were lowest at the beginning of the interval cycles, increased as the rate of responding increased, and then decreased during the timeout periods. At all parameters studied, there was a direct relationship between changes in response rate within fixed‐interval cycles and changes in heart rate and blood pressure. The results suggest that a much closer concordance of these cardiovascular parameters and schedule‐controlled responding is obtained by examining ongoing behavior as it occurs within the contingencies by which it is maintained.