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.     

Relationship between response rate and reinforcement frequency in variable-interval schedules: III. The effect of d-amphetamine.

Four rats were exposed to variable-interval schedules specifying a range of different reinforcement frequencies. The effects of two doses of d-amphetamine (1.6 and 3.2 mumol/kg) upon performance maintained under each schedule were examined. In the case of each rat, the response rates observed under control conditions (no injection or injection of the vehicle alone) were increasing, negatively accelerated functions of reinforcement frequency, the data conforming closely to Herrnstein's (1970) equation. In each rat, d-amphetamine (3.2 mumol/kg) significantly reduced the value of the constant Rmax, which expresses the theoretical maximum response rate. In each rat, the value of KH, which expresses the reinforcement frequency needed to obtain the half-maximal response rate, was also reduced, although this only achieved statistical significance in the case of one rat. When the proportional change in response rate in the presence of the drug was plotted against the response rate under control conditions on double logarithmic co-ordinates, linear functions of negative slope were obtained; in each rat the slope was steeper and the value of the control response rate at which d-amphetamine exerted no effect was lower in the case of the higher dose (3.2 mumol/kg) than in the case of the lower dose (1.6 mumol/kg).     

Effects of d-amphetamine, cocaine, and phencyclidine on the acquisition of response sequences with and without stimulus fading.

In each of three components of a multiple schedule, monkeys were required to emit a different sequence of four responses in a predetermined order on four levers. Sequence completions produced food on a fixed-ratio schedule. Errors produced a brief timeout. One component of the multiple schedule was a repeated-acquisition task where the four-response sequence changed each session (learning). The second component of the multiple schedule was also a repeated-acquisition task, but acquisition was supported through the use of a stimulus-fading procedure (faded learning). In a third component of the multiple schedule, the sequence of responses remained the same from session to session (performance). At higher doses, d-amphetamine, cocaine, and phencyclidine decreased the overall rate of responding and increased the percent errors in all three components. At lower doses, however, the three drugs produced selective effects on errors. Errors were increased in the learning component at lower doses than those required to disrupt the behavior in the faded-learning component. The performance component tended to be the least sensitive to disruptive drug effects. The data are consistent with the view that stimulus fading can modulate the effects of drugs on acquisition.     

The effects of reinforcement frequency and response requirements on the maintenance of behavior.

Six rats responded under fixed-interval and tandem fixed-interval fixed-ratio schedules of food reinforcement. Basic fixed-interval schedules alternated over experimental conditions with tandem fixed-interval fixed-ratio schedules with the same fixed-interval value. Fixed-interval length was varied within subjects over pairs of experimental conditions; the ratio requirement of the tandem schedules was varied across subjects. For both subjects with a ratio requirement of 10, overall response rates and running response rates typically were higher under the tandem schedules than under the corresponding basic fixed-interval schedules. For all subjects with ratio requirements of 30 or 60, overall response rates and running response rates were higher under the tandem schedules than under the corresponding basic fixed-interval schedules only with relatively short fixed intervals. At longer fixed intervals, higher overall response rates and running rates were maintained by the basic fixed-interval schedules than by the tandem schedules. These findings support Zeiler and Buchman's (1979) reinforcement-theory account of response strength as an increasing monotonic function of both the response requirement and reinforcement frequency. Small response requirements added in tandem to fixed-interval schedules have little effect on reinforcement frequency and so their net effect is to enhance responding. Larger response requirements reduce reinforcement frequency more substantially; therefore their net effect depends on the length of the fixed interval, which limits overall reinforcement frequency. At the longest fixed intervals studied in the present experiment, reinforcement frequency under the tandem schedules was sufficiently low that responding weakened or ceased altogether.     

d-amphetamine and fixed-interval performance: effects of operant history.

Sixteen rats were initially exposed for 50 sessions to either a fixed-ratio 40 or an interresponse-time-greater-than-11-second food reinforcement schedule, then shifted to a fixed-interval 15-second food reinforcement schedule. Animals with fixed-ratio 40 histories lever pressed at much higher rates under the fixed-interval schedule than did animals with inter-response-time-greater-than-11-second histories. This difference persisted across 93 sessions of fixed-interval exposure. The effects of d=amphetamine were assessed after 15 and 59 sessions of fixed-interval exposure. On both occasions, the low-rate responding of animals with interresponse-time-greater-than-11-second histories was typically increased by all doses of the drug, while the high-rate responding of animals with fixed-ratio 40 histories was typically decreased by all doses of the drug. These results suggest that control response rate under the fixed-interval schedule, which may be affected by a history of responding under another schedule, is the primary determinant of the relative effects of d-amphetamine.     

Conditioned reinforcement dynamics in three-link chained schedules.

In two experiments rats were trained on three-link concurrent-chains schedules of reinforcement. In Experiment 1, additional entries to one terminal link were added during one of the middle links to a baseline schedule that was otherwise equal for the two chains, and, depending on the condition, these additional terminal-link presentations ended either in food or in no food. When food occurred, preference was always in favor of the chain with the additional terminal-link presentations (which also entailed a higher rate of reinforcement). When no food occurred at the end of the additional terminal links, the outcome depended on the nature of the stimuli associated with these additional terminal links. When stimuli different from the reinforced baseline terminal links were used for the no-food terminal links, preference was against the choice alternative that led to the extra periods of extinction. When the same stimulus was used for the two kinds of terminal links, preference was near indifference, that is, significantly greater than when different stimuli were used. In Experiment 2, rats learned repeated reversals of a simultaneous discrimination under a three-link concurrent-chains schedule, in which the food or no-food choice outcomes were delayed until the end of the chain. Different conditions were defined by the point in the chain at which differential stimuli occurred. When the middle and terminal links provided no differential stimuli, discrimination was acquired more slowly than when differential stimuli occurred in both links. When differential stimuli occurred in the middle but not the terminal links, acquisition rates were intermediate. Both experiments together show that the effects of stimuli in a chain schedule are due partly to the time to food correlated with the stimuli and partly to the time to the next conditioned reinforcer in the sequence.     

Molecular analyses of the effects of d-amphetamine on fixed-interval schedule performances of rats.

A series of doses (0.5 to 2.0 mg/kg) of d-amphetamine was administered to rats whose lever pressing was maintained by fixed-interval 30-s, 60-s, or 120-s schedules of reinforcement by sucrose delivery. Under both saline and d-amphetamine conditions, molecular features of responding were reliably described in terms of the distribution of postreinforcement pauses and local response rate following the onset of responding. Postreinforcement pause always varied from interval to interval but, on average, shortened under the drug. Local response rate (response rate exclusive of pause time) tended to decrease under the drug, and where acceleration occurred within runs of responses, it was reduced by the drug. All of these effects were dose-related. These findings suggest that fixed-interval behavior can be analyzed effectively at a molecular level, and that the effects of d-amphetamine are best described as disruption of temporal discrimination.     

Effects of D-amphetamine on response acquisition with immediate and delayed reinforcement.

The present study examined in 8-hour sessions the effects of d-amphetamine (1.0, 5.6, and 10 mg/kg) on the acquisition of lever-press responding in rats that were exposed to procedures in which water delivery was delayed by 0, 8, or 16 seconds relative to the response that produced it. Both nonresetting- and resetting-delay conditions were studied. Although neither shaping nor autoshaping occurred, substantial levels of operative-lever responding developed under all conditions in which responses produced water. The lowest dose (1.0 mg/kg) of d-amphetamine either had no effect on or increased operative-lever pressing, whereas higher doses typically produced an initial reduction in lever pressing. Nonetheless, overall rates of operative-lever pressing at these doses were as high as, or higher than, those observed with vehicle. Thus, response acquisition was observed under all reinforcement procedures at all drug doses. In the absence of the drug, most responding occurred on the operative lever when reinforcement was immediate. Such differential responding also developed under both nonresetting- and resetting-delay procedures when the delay was 8 seconds, but not when it was 16 seconds. d-Amphetamine did not affect the development of differential responding under any procedure. Thus, consistent with d-amphetamine's effects under repeated acquisition procedures, the drug had no detrimental effect on learning until doses that produced general behavioral disruption were administered.     

Control over response number by a targeted percentile schedule: reinforcement loss and the acute effects of d-amphetamine.

Two fixed-consecutive-number-like procedures were used to examine effects of acute d-amphetamine administration on control over response number. In both procedures, rats were required to press the left lever at least once and then press the right lever to complete a trial. The consecutive left-lever presses on each trial comprised a "run." Under the targeted percentile schedule, reinforcement was provided if the current run length was closer to the target length (16) than half of the most recent 24 runs. This differentially reinforced run length while holding reinforcement probability constant at .5. A second group acquired the differentiation under the targeted percentile schedule, but were then shifted to a procedure that yoked reinforcement probability by subject and run length to that obtained under the targeted percentile schedule. The two procedures generated practically identical control run lengths, response rates, reinforcement probabilities, and reinforcement rates. Administration of d-amphetamine disrupted percentile responding to a greater degree than yoked control responding. This disruption decreased reinforcement frequency less in the former than the latter procedure. The similar baseline responding under these two procedures suggests that this difference in sensitivity was due to behavioral adjustments to drug prompted by reduction of reinforcement density in the yoked control but not the percentile schedule. These adjustments attenuate the drug's effects under the former, but not the latter, procedure.     

Second-order schedules of token reinforcement: effects of varying the schedule of food presentation

In the initial link of a complex schedule, one discriminative stimulus was presented and lever pressing produced tokens on fixed-ratio schedules. In the terminal link, signalled by a second discriminative stimulus, deposits of the tokens produced food. With two rats, the terminal link was presented after each sixth component schedule of token reinforcement was completed. With the other two rats, the terminal link was presented following the first component schedule completed after a fixed interval. During the terminal link, each token deposit initially produced food. The schedule of food presentation was subsequently increased such that an increasing number of token deposits in the terminal link was required for each food presentation. Rates of lever pressing in the initial link were inversely related to the schedule of food presentation in the terminal link. These results are similar to those of experiments that have varied schedules of food presentation in chained schedules. Rates and patterns of responding controlled throughout the initial link were more similar to those ordinarily controlled by second-order brief-stimulus schedules than to those controlled by comparable extended chained schedules.     

Body weight and response acquisition with delayed reinforcement.

The relation between body weight and responding established with unsignaled delayed reinforcement was investigated. In three experiments, naive rats were deprived to either 70%, 80%, or 90% of ad libitum weight and were then exposed to tandem variable-interval 15-s differential-reinforcement-of-other-behavior 30-s schedules. The tandem schedule defined a resetting unsignaled delay-of-reinforcement procedure. In the first experiment, speed of magazine training, acquisition of lever pressing, and final rate of lever pressing were related to body weight. In the next experiment, lever pressing was established and maintained in rats that were magazine trained at 70% of ad libitum weight but that were then exposed to the delay procedure at 90% of ad libitum weight. Responding did not change consistently either across or within subjects in subsequent conditions in which body weight was manipulated. In the final experiment, lever pressing was established and maintained with delayed reinforcement in the absence of magazine training for each of 2 rats at 70% and for 1 of 2 rats at 90% of ad libitum weight. The results further illuminate the conditions under which responding can be established in the absence of training and when such responses are reinforced only following an unsignaled delay period.     

Variable-ratio schedules of timeout from avoidance: effects of d-amphetamine and morphine.

Rats were trained on concurrent schedules in which pressing one lever postponed shock and pressing the other lever produced periods of signaled timeout from avoidance on variable-ratio schedules. These procedures generated high rates of timeout-reinforced responding and provided a baseline for studying the effects of drugs on behavior maintained by different types of negative reinforcement (shock postponement vs. timeout). Morphine (2.5 to 10.0 mg/kg) reduced behavior maintained by timeout at doses that increased or had no effect on avoidance responding. In contrast, d-amphetamine (0.125 to 2.0 mg/kg) produced large increases in timeout responding at doses that had minimal effect on avoidance in rats trained on variable-interval and variable-ratio schedules. Thus, the event-dependent effects of morphine, observed in previous studies in which timeout responding was maintained at low rates by interval schedules, were replicated with high timeout rates maintained by variable-ratio schedules. The effects of d-amphetamine could also be described as "event dependent" because timeout responding was stimulated more than avoidance regardless of the maintenance schedule or baseline rate.     

The control of responding by sounds: unusual effect of reinforcement.

Naive rats were trained to respond on one lever in the presence of noise bursts from one speaker and on a second lever in the presence of noise bursts from a second speaker. The speakers were mounted behind the levers. When responding on the lever adjacent to the sounding speaker was reinforced, control developed within fewer than five trials. When responding on the nonadjacent lever was selectively reinforced, responding on the lever adjacent to the sounding speaker increased in probability for several sessions. Naive rats were trained to respond on the nonadjacent lever following preexposure to the sound. Responding on the lever adjacent to the sounding speaker increased in probability, showing that novelty was not responsible for the effect. Naive rats were run on automaintenance procedures in which there was no explicit pairing of sound and magazine operation, 100% pairing of sound and magazine operation, or magazine operation following 40% of sound presentations. None of the rats acquired the response of approaching and sniffing the sounding speaker, indicating that sound-magazine pairing was not responsible for the effect.     

Contingency and stimulus change in chained schedules of reinforcement

Higher rates of pecking were maintained by pigeons in the middle component of three-component chained fixed-interval schedules than in that component of corresponding multiple schedules (two extinction components followed by a fixed-interval component). This rate difference did not occur in equivalent tandem and mixed schedules, in which a single stimulus was correlated with the three components. The higher rates in components of chained schedules demonstrate a reinforcing effect of the stimulus correlated with the next component; the acquired functions of this stimulus make the vocabulary of conditioned reinforcement appropriate. Problems in defining conditioned reinforcement arise not from difficulties in demonstrating reinforcing effects but from disagreements about which experimental operations allow such reinforcing effects to be called conditioned.     

Drug discrimination: stimulus control during repeated testing in extinction

Rats were trained, under a two-lever drug-discrimination procedure, to respond differentially depending upon whether lorazepam (1.0 mg/kg) or no injection had been administered before the session. Responses on the appropriate lever produced a food pellet under a modified fixed-ratio (FR) 10 schedule, in which the 10 responses had to be emitted consecutively. In reinforcement tests, completing an FR 10 on either lever produced a pellet. In extinction tests, stimulus changes paired with reinforcement occurred but no pellet was delivered. Training sessions were conducted between test sessions. Each of four extinction phases consisted of six tests preceded by one stimulus (e.g., lorazepam). Repeated exposures to extinction reduced response rates for all rats, but stimulus control, as inferred from either percentage of total responses or percentage of total FR 10s on the drug-appropriate lever, remained high. The percentage of total FR 10s measure was less subject to skewing under low-rate conditions than was the percentage of total responses measure and provided an evaluation of stimulus control in terms of meeting the consecutive response contingency. These results demonstrate a level of independence between response rate and stimulus control in drug discrimination, which has positive implications for the validity of interpreting discriminative effects of novel test conditions in well-trained animals, even when overall response rates are low.     

Behavior of rats under fixed consecutive number schedules: effects of drugs of abuse.

Four rats responded under a simple fixed consecutive number schedule in which eight or more consecutive responses on the run lever, followed by a single response on the reinforcement lever, produced the food reinforcer. Under this simple schedule, dose-response curves were determined for diazepam, morphine, pentobarbital, and phencyclidine. The rats were then trained to respond under a multiple fixed consecutive number schedule in which a discriminative stimulus signaled when the response requirement on the run lever had been completed in one of the two fixed consecutive number component schedules. Under control conditions, the percentage of reinforced runs under the multiple-schedule component with the discriminative stimulus added was much higher than the percentage of reinforced runs under the multiple-schedule component without the discriminative stimulus. All of the drugs decreased the percentage of reinforced runs under each of the fixed consecutive number schedules by increasing the conditional probability of short run lengths. This effect was most consistently produced by morphine. The drugs produced few differences in responding between the multiple fixed consecutive number components. Responding under the simple fixed consecutive number schedule, however, was affected at lower doses of the drugs than was responding under the same fixed consecutive number schedule when it was a component of the multiple schedule. This result may be due to the difference in schedule context or, perhaps, to the order of the experiments.     

An experimental analysis of the effects of d-amphetamine and cocaine on the acquisition and performance of response chains in monkeys.

In one component of a multiple schedule of food presentation, monkeys acquired a different four-response chain each session by responding sequentially on three keys in the presence of four geometric forms (learning). In the other component, the four-response chain was the same each session (performance). Both d-amphetamine and cocaine, at the higher doses, disrupted the behavior in the learning component; the overall response rate decreased, the overall accuracy was impaired (i.e., percent errors increased), and there was less within-session error reduction. The performance component was generally less sensitive than the learning component to the disruptive effects of both drugs on rate and accuracy. After pre-feeding or during an extended session, the response rate decreased in both components, but accuracy was generally unaffected. When the four discriminative stimuli in both components were removed, the behavior was disrupted to a greater extent in the performance component. The disruptive effects of both drugs on behavior in the learning component were attenuated when the drugs were administered during the session after the response chain had been acquired. It was concluded that the greater sensitivity of the learning component to disruptive drug effects is related to the relatively weak stimulus control and/or the lower rate of reinforcement associated with that component.     

Effects of schedule of reinforcement on a pentobarbital discrimination in rats.

The purpose of this study was to determine the effects of the schedule of reinforcement on a pentobarbital discrimination in rats. Five rats were trained to discriminate 10 mg/kg pentobarbital from saline under a multiple fixed-interval 180-s fixed-ratio 20 schedule of reinforcement. During both saline and pentobarbital training sessions, subjects emitted a higher percentage of correct responses under the fixed-ratio component as compared to the fixed-interval component of the multiple schedule. Determination of the pentobarbital dose-response curve under the fixed-ratio component resulted in a steep curve characterized by responding on the saline lever at low doses and on the drug lever at higher doses. Under the fixed-interval component, a graded dose-effect curve was produced, with considerable responding on both levers after intermediate doses of pentobarbital. The administration of phencyclidine and MK-801 resulted in an intermediate level of drug-lever responding for some subjects. Administration of d-amphetamine resulted in saline (nondrug) appropriate responding. The results of this study demonstrate that the schedule of reinforcement is a determinant of drug stimulus control, just as it is a determinant of other drug effects.     

Behavioral effects of delayed timeouts from reinforcement

Timeouts are sometimes used in applied settings to reduce target responses, and in some circumstances delays are unavoidably imposed between the onset of a timeout and the offset of the response that produces it. The present study examined the effects of signaled and unsignaled timeouts in rats exposed to concurrent fixed‐ratio 1 fixed‐ratio 1 schedules of food delivery, where each response on one lever, the location of which changed across conditions, produced both food and a delayed 10‐s timeout. Delays of 0 to 38 s were examined. Delayed timeouts often, but not always, substantially reduced the number of responses emitted on the lever that produced timeouts relative to the number emitted on the lever that did not produce timeouts. In general, greater sensitivity was observed to delayed timeouts when they were signaled. These results demonstrate that delayed timeouts, like other delayed consequences, can affect behavior, albeit less strongly than immediate consequences.     

Effects of magnitude of food reinforcement on free-operant response rates

In Experiment 1 rats were trained to press a lever on a variable-ratio schedule of food presentation and were then exposed to progressively increasing magnitudes of food reinforcement. Response running rates (rates exclusive of the postreinforcement pause) were found to increase as a function of increasing reinforcement magnitudes. The effect of reinforcement magnitude on response rates inclusive of the postreinforcement pause, however, was less pronounced. Increases in the magnitude of reinforcement were also found to increase the length of the postreinforcement pause. Rats in Experiment 2 were trained to respond on a chained differential-reinforcement-of-low-rate variable-ratio schedule, and were exposed to increasing magnitudes of reinforcement as in Experiment 1. Response running rates increased in the variable-ratio component but decreased in the other component of the schedule. The results are discussed with reference to incentive accounts of reinforcement and the action of reinforcement on the response units generated by the operative contingencies.