The contribution of an added counter to a fixed-ratio schedule

Although previous research showed that a visual counter increased the rate of responding on a large fixed-ratio schedule, a theoretical analysis of the factors responsible for fixed-ratio performance suggests that the primary control by number of responses since reinforcement is to weaken the performance. The present experiment employed a multiple schedule in which the same fixed-ratio value alternated with and without an added counter. It tested the hypothesis that the differential reinforcement of high-rate responding masked the attenuation of the fixed-ratio performance from the unoptimal discriminative control produced by the fixed relation between number of responses and reinforcement. In the present experiment the postreinforcement pause was consistently longer in the components with the added counter, while running rates remained comparable between the components of the multiple schedule. Both components of the multiple schedule involved differential reinforcement of high-rate responding while only the components with the added counter amplified the discriminative control by number of pecks since reinforcement.     

Drug effects on repeated acquisition: comparison of cumulative and non-cumulative dosing

Pigeons acquired a different four-response chain each session by responding sequentially on three keys in the presence of a sequence of four colors. The response chain was maintained by food presentation under a fixed-ratio schedule. Errors produced a brief timeout but did not reset the chain. Each day there were four 15-minute sessions, with a 10-minute inter-session interval. Cumulative dose-effect curves for phencyclidine, pentobarbital, and d-amphetamine were obtained by giving an injection before each of the four sessions; successive injections increased the cumulative dose in equally spaced logarithmic steps. For comparison, non-cumulative doses of each drug (i.e., doses not preceded by other doses on the same day) were also tested. As the cumulative dose of each drug increased, the overall response rate decreased, the percent errors increased, and there was less within-session error reduction (acquisition). With phencyclidine and pentobarbital, the rate-decreasing and error-increasing effects tended to be greater with a non-cumulative dose than with the corresponding cumulative dose. In contrast, with d-amphetamine, the effects were considerably greater with the cumulative doses. The results indicate that although the cumulative-dosing procedure saved a substantial amount of time in determining dose-effect curves, there were quantitative differences in effects between cumulative and non-cumulative doses.     

Drug discrimination under a concurrent fixed-ratio fixed-ratio schedule.

Pigeons were trained to discriminate 5.0 mg/kg pentobarbital from saline under a two-key concurrent fixed-ratio 10 fixed-ratio 40 schedule of food presentation, in which the fixed-ratio component with the lower response requirement was programmed to reinforce responding on one key after drug administration (pentobarbital-biased key) and on the other key after saline administration (saline-biased key). After responding stabilized, pigeons averaged 98% of their responses on the pentobarbital-biased key during training sessions preceded by pentobarbital, and they averaged 90% of their responses on the saline-biased key during training sessions preceded by saline. In test sessions preceded by doses of pentobarbital, chlordiazepoxide, or ethanol, pigeons switched from responding on the saline-biased key at low doses to responding on the pentobarbital-biased key at higher doses (the dose-response curve was quantal). High doses of phencyclidine produced responding on both keys, whereas pigeons responded almost exclusively on the saline-biased key after all doses of methamphetamine. These and previous experiments using concurrent reinforcement schedules to study drug discrimination illustrate that the schedule of reinforcement is an important determinant of the shape of dose-effect curves in drug-discrimination experiments.     

Cocaine and food deprivation: effects on food-reinforced fixed-ratio performance in pigeons.

Key pecking by 6 pigeons was maintained by a fixed-ratio 30 schedule of food presentation while body weights were 80% of free-feeding weights. Acute administration of cocaine (0.3 to 13.0 mg/kg, i.m.) dose-dependently decreased response rates. Dose-effect curves were shifted to the right when 3 of the 6 pigeons were maintained at 70% of free-feeding weights and were shifted to the left when the other 3 pigeons were maintained at 90% of free-feeding weights. Then a dose of cocaine that initially decreased response rates by more than 95% of control rates was administered before each daily session. Comparable degrees of tolerance to these rate-decreasing effects developed in the two groups. The rate at which responding recovered was relatively rapid for pigeons in the 70% free-feeding-weight group and was slower for 2 of the 3 pigeons in the 90% free-feeding-weight group. When body weights were then increased from 70% to 80% or were decreased from 90% to 80% of free-feeding weight, performance was disrupted initially only for pigeons whose weight went from 70% to 80% of free feeding. In the present experiment the degree of deprivation may have indirectly influenced the degree of tolerance that developed to cocaine's response rate-decreasing effects because it directly influenced the dose chosen to be administered chronically. The degree of deprivation appeared to have a more direct influence on the rate at which tolerance developed.     

Bias of phencyclidine discrimination by the schedule of reinforcement.

Pigeons, trained to discriminate phencyclidine from saline under a procedure requiring the bird to track the location of a color, received cumulative doses of phencyclidine, pentobarbital, or d-amphetamine with a variety of schedules of reinforcement in effect (across phases). When the same second-order schedules were used to reinforce responding after either saline or phencyclidine administration, stimulus control by phencyclidine did not depend on the schedule parameter. When different second-order schedules were used that biased responding toward the phencyclidine-correlated key color, pigeons responded on the phencyclidine-correlated key at lower doses of phencyclidine and pentobarbital than when the second-order schedule biased responding toward the saline key color. A similar but less marked effect was obtained with d-amphetamine. Attempts to produce bias by changing reinforcement magnitude (duration of food availability) were less successful. A signal-detection analysis of dose-effect curves for phencyclidine under two of the second-order schedules employed suggested that at low doses of phencyclidine, response bias is a major determinant of responding. As doses were increased, position preferences occurred and response bias decreased; at higher doses both response bias and position preference decreased and discriminability increased. With low doses of pentobarbital, responding again was biased but increasing doses produced position preference with only small increases in discriminability. At low doses of d-amphetamine responding also was biased, but bias did not decrease consistently with dose nor did discriminability increase. These experiments suggest that the schedule of reinforcement can be used to bias responding toward or away from making the drug-correlated response in drug discrimination experiments, and that signal-detection analysis and analysis of responding at a position can be used to separate the discriminability of the drug state from other effects of the drug on responding.     

Drug effects on responding maintained by stimulus-reinforcer and response-reinforcer contingencies.

The effects of pentobarbital and d-amphetamine were assessed on key pecking by pigeons under conventional single-key multiple schedules and under two-key multiple schedules in which discriminative stimuli appeared on one key (stimulus key) while pecks on a second key (constant key) produced food. Pecks on the stimulus key had no scheduled consequences. A 60-second variable-interval schedule operated in one component of each multiple schedule: either extinction or a 60-second variable-time schedule operated in the alternate component. When the alternate-component schedule was extinction, a high rate of responding was maintained in the variable-interval component of the single-key schedule; responding on both keys was maintained in the variable-interval component of the two-key schedule. Pentobarbital increased responding in the variable-interval component of the single-key schedule and increased stimulus-key, but not constant-key responding in that component of the two-key schedule. When the alternate-component schedule was changed to variable time, responding declined in the variable-interval component of the single-key schedule; stimulus-key responding was no longer maintained under the two-key schedule. Pentobarbital decreased responding in the variable-interval component of both schedules. With an exception, d-amphetamine only decreased responding in the variable-interval component of the single- and two-key schedules both when the alternate-component schedule was extinction and when it was variable time. The results suggest that the effects of pentobarbital, but not d-amphetamine, depend on the nature of the contingency (stimulus-reinforcer, response-reinforcer) that maintains responding.     

Effects of d-amphetamine and pentobarbital under concurrent fixed-ratio schedules.

Pigeons were studied under a two-key concurrent fixed-ratio schedule of food presentation. During the first five sessions, the fixed-ratio requirements were 30 responses on one key (major key) and 120 responses on the other key (minor key): responding occurred almost exclusively on the major key. When the fixed-ratio requirements were then made equal at 30 responses on both keys, responding continued to predominate on the major key. The asymmetric distribution of responses persisted when the concurrent fixed-ratio fixed-ratio schedule was interrupted with periods during which the major key was associated with extinction while the other key remained associated with a fixed-ratio schedule. Additionally, in some subjects the fixed-ratio requirements were increased. These schedule modifications decreased the asymmetry in responding but did not eliminate it. d-Amphetamine decreased rates on both keys and slightly increased the asymmetric distribution of responses, while pentobarbital reversed the distribution of responses by increasing low rates and decreasing high rates. The pigeons maintained their original asymmetric distribution of responses during the 1 1/2-year-long study, despite schedule alterations and drug administrations.     

Mirror pecking and timeout under a multiple fixed-ratio schedule of food delivery

Pigeons were trained to peck a key under a multiple fixed-ratio 25 fixed-ratio 175 schedule of food presentation. In the first condition, either a mirror or the opportunity to produce a 30-second timeout were available. In a second condition, mirror and timeout availability were reversed for the two groups. Following a return to the initial condition, mirror and timeout keys were presented together for all birds. Mirror and timeout responses occurred predominantly in the pause in the larger fixed-ratio component, regardless of whether the opportunities for the two responses were available singly or together. Mirror responding occurred in a greater proportion of the pauses than did timeouts. When the opportunities for both mirror pecking and timeout were available concurrently, they occurred with probabilities similar to those under the single conditions. Within the pause itself, mirror responses most frequently occurred immediately after reinforcement. Timeouts occurred most frequently toward the end of the pause, and some timeouts occurred in the early part of the run. Longer preratio pausing occurred in the larger fixed-ratio component in the conditions in which the mirror was present, whether or not any mirror pecks were recorded.     

Drug discrimination under a concurrent schedule.

Three pigeons were trained to discriminate a 5.0 mg/kg dose of pentobarbital from saline under a two-key concurrent schedule with responding on the key associated with the presession injection, under both stimulus conditions, producing four times as many reinforcers as responding on the other key. This concurrent schedule resulted in approximately 70% responding to the higher reinforcement key under the pentobarbital stimulus and approximately 30% responding to that key under the saline stimulus. During testing, then, the pigeons were able to dose-dependently emit higher (>70%) or lower (<30%) values than were established under the control conditions. Dose-response curves were determined for pentobarbital (twice), methamphetamine, phencyclidine, chlordiazepoxide, and the combination of pentobarbital and the barbiturate antagonist bemegride. The results obtained with pentobarbital and chlordiazepoxide showed that, as the dose increased, pentobarbital-appropriate responding also increased. Methamphetamine produced relatively flat dose--response curves, whereas phencyclidine administration produced inconsistent effects on responding. The combination of the training dose of pentobarbital with increasing doses of bemegride produced a decrease in pentobarbital-appropriate responding. The results also showed that the dose-response curves for pentobarbital and chlordiazepoxide, instead of being all or none, were graded functions of the drug dose.     

The effects of brief-stimulus presentations in fixed-ratio second-order schedules

Pigeons' responses were reinforced according to a three-component multiple schedule. In Component 1, key pecks produced food according to a fixed-ratio second-order schedule with fixed-ratio units. Here, a fixed number of fixed-ratio units produced food, and the brief stimulus terminating each unit also accompanied food. Responses in Component 2 produced food on an identical schedule except that the brief stimulus was not paired with food. Component 3 contained a simple fixed-ratio schedule whose response requirement equaled that of Components 1 and 2. Across conditions the size of the fixed-ratio unit (five, ten, twenty, forty, and eighty responses) and the total number of responses per reinforcement were parametrically manipulated. The highest response rates and shortest preratio pauses were observed in Component 3 (no brief stimulus). The lowest rates and longest pauses were found in the component with paired brief-stimulus presentations, indicating that the food-paired brief stimulus suppressed responding. The suppressive effects were greatest when the fixed-ratio units were small (e.g., fixed-ratio 5) and the total fixed-ratio requirement was large (e.g., fixed-ratio 160). Under no conditions did the paired brief stimulus facilitate responding. The nonpaired brief stimulus also suppressed responding but to a lesser extent. The suppressive effects of nonpaired brief stimuli were greatest when the fixed-ratio units were small and the total response requirement was large. These data suggest that the suppressive effects of the brief stimuli may have masked the conditioned-reinforcing effects reported in other studies, and that conditions that maximize suppression in second-order schedules involve the use of fixed-ratio schedule units and the presentation of many brief stimuli per reinforcer.     

Effects of d-amphetamine and cocaine on strained ratio behavior in a repeated-acquisition task.

Pigeons acquired a different four-response chain each session by responding sequentially on three keys in the presence of four colors. When the fixed-ratio requirement for food presentation was five completions of the chain, d-amphetamine and cocaine disrupted the behavior. As the dose of each drug was increased, the overall response rate decreased, the overall accuracy was impaired (i,e., percent errors increased), and there was less within-session error reduction (acquisition). In contrast, when the fixed-ratio requirement was either 20 or 50 completions of the chain, certain doses of both drugs produced large increases in the overall response rate by eliminating the extended pausing (ratio strain) that was characteristic of the control sessions. These rate-increasing effects were accompanied by error-decreasing effects, both during acquisition and after the response chain had been acquired. Taken together, the results show that the effects of d-amphetamine and cocaine on behavior in a repeated-acquisition task can be modulated by manipulating the value of the fixed-ratio schedule maintaining the behavior.     

Neurochemical changes correlated with behavior maintained under fixed-interval and fixed-ratio schedules of reinforcement.

Key pecking of 4 pigeons was maintained under a multiple 3-min fixed-interval, 30-response fixed-ratio schedule of food presentation. Only one schedule was in effect during an experimental session, and each was correlated with a different keylight stimulus and location (left vs. right). The different schedule components alternated across days or weeks. Cerebrospinal fluid was collected from chronically implanted intracerebroventricular cannulae following sessions with the different schedules, as well as following sessions in which reinforcement was withheld (extinction), when response-independent food was delivered, and when the experimental chamber was dark and there were no scheduled events. Metabolites of the neurotransmitters serotonin, norepinephrine, and dopamine were assayed in cerebrospinal fluid using high-performance liquid chromatography with electrochemical detection. Compared to the fixed-ratio condition, responding maintained under the fixed-interval schedule resulted in consistently higher levels of the serotonin metabolite 5-hydroxyindoleacetic acid and of the dopamine metabolite homovanillic acid in all pigeons. Levels of 3-methoxy-4-hydroxyphenylethylene glycol, a metabolite of norepinephrine, and dihydroxyphenylacetic acid, another dopamine metabolite, were also higher in 3 of the 4 pigeons following exposure to the fixed-interval schedules when compared to levels of these metabolites after exposure to the fixed-ratio schedule. Extinction of fixed-ratio responding resulted in large increases in 5-hydroxyindoleacetic acid compared to levels of this metabolite under the fixed-ratio schedule, whereas this serotonin metabolite decreased during extinction of responding under the fixed-interval schedule. Control procedures suggested that the neurochemical changes were not related to the rate of responding but were a function of the specific experimental conditions. Distinctive neurochemical changes that accompany schedule-controlled responding show the sensitivity of the neurochemical environment to behavioral contingencies and demonstrate further the profound impact that such contingencies have on biobehavioral processes.     

Selective antagonism of the rate-decreasing effect of d-amphetamine by chlorpromazine in a repeated-acquisition task.

Pigeons acquired a different four-response chain each session by responding sequentially on three keys in the presence of four colors. The response chain was maintained by food presentation under a fixed-ratio schedule. When d-amphetamine was administered alone, the overall response rate decreased and the percent errors increased with increasing doses. When a small dose of chlorpromazine, which was ineffective when given alone, was administered in combination with d-amphetamine, the rate-decreasing effect was antagonized. The antagonism was selective, however, in that the error-increasing effect of d-amphetamine was augmented by chlorpromazine. The nature of the joint effect of the two drugs thus depended on the behavioral measure: rate vs. accuracy.     

Reinforcing the absence of fixed-ratio performance

Pigeons received food for key pecking according to a fixed-ratio schedule, while, at the same time, food also was available for not pecking for a specified time. With a fixed ratio of 60, responding was not affected by not-pecking times of 80 or 40 seconds, and was eliminated completely at 10 seconds. With ratios of 180, pecking stopped with not-pecking times of 80 seconds or less; with ratios of 300, it stopped at 120 seconds or less. Not-responding schedules produced steady-state performance immediately following contact with the schedule. With return to the fixed-ratio schedule alone, response rate sometimes was elevated temporarily. When response-independent food presentations replaced the not-pecking schedule, response rate often was enhanced, and the ratio pattern was lost. Only the highest densities of food delivery eliminated responding, even with a fixed ratio of 300. In general, the effects corresponded to those of punishment, except that contrast had appeared both during and after punishment, and now appeared only after the response elimination procedure was suspended.     

Effects of d-amphetamine on responding under second-order schedules of reinforcement with paired and nonpaired brief stimuli.

Three pigeons were studied under a multiple schedule in which pecks in each component were reinforced according to a variable-interval 120-s second-order schedule with fixed-interval 60-s units. In the first component of the multiple schedule, the completion of a fixed interval produced either food or a 4-s change in key color plus houselight illumination. In the second component an identical schedule was in effect, but the stimulus was a 0.3-s change in key color. Both long and short brief stimuli were not paired with food presentations in Conditions 1 and 3 and were paired with food in Condition 2. There were no consistent differences in response patterns under paired and nonpaired brief-stimulus conditions when the stimulus was a 4-s change in key color accompanied by houselight illumination. However, pairing the 0.3-s key-color change with food presentations resulted in higher indices of curvature and lower response rates in the early segments of the fixed interval than when the stimulus was not paired with food presentations. Low doses of d-amphetamine (0.3 and 1 mg/kg) produced small and inconsistent increases in overall response rates, and higher doses (3 and 10 mg/kg) decreased overall response rates. d-Amphetamine altered response patterns within fixed intervals by decreasing the indices of curvature and increasing response rates in the early segments of the fixed interval. Response rates and patterns under paired and nonpaired brief-stimulus conditions were not differentially affected by d-amphetamine. Thus, evidence for the enhancement of the conditioned reinforcement effects of psychomotor stimulant drugs was not found with the second-order schedules used in the present study.     

Effects of ethanol on multiple fixed-interval fixed-ratio schedule performances: dynamic interactions at different fixed-ratio values.

Key pecking by three pigeons was maintained under a multiple fixed-interval fixed-ratio schedule of food presentation. The fixed-interval value remained at 3 minutes, while the fixed-ratio size was increased systematically in 30-response increments from 30 to either 120 (two pigeons) or 150 (one pigeon). At least two lower fixed-ratio values were also redetermined. The effects of ethanol (5 to 2.5 g/kg) were assessed at each of the different schedule parameters. Both overall and running response rates under the fixed-ratio schedule decreased with increases in the size of the fixed-ratio schedule; pause duration under the fixed-ratio schedule was directly related to increases in fixed-ratio size. Overall and running rates of responding under the fixed-interval schedule changed little with increases in the size of the fixed-ratio schedule. Despite the relative invariance of fixed-interval responding across the different fixed-ratio values, the effects of ethanol on responding under the fixed-interval schedule differed depending on the size of the fixed-ratio schedule. Greater increases occurred in both overall and in lower local rates of responding under the fixed-interval schedule when the fixed-ratio value was 120 or 150. The effects of ethanol on responding under the fixed-ratio schedule also depended on the size of the fixed ratio. Increases in responding under the fixed-ratio schedule were typically greater at the higher fixed-ratio values where response rates were lower. When the effects of ethanol were redetermined at the lower fixed-ratio parameter values, rates and patterns of responding were comparable to those obtained initially. However, the dose-effect curves for responding under both fixed-ratio and fixed-interval schedules were shifted up and to the right of those determined during the ascending series. The effects of ethanol can depend on rate or responding, behavioral history, and the context in which behavior occurs.     

Tolerance to effects of cocaine on behavior under a response-initiated fixed-interval schedule

Tolerance to effects of cocaine can be modulated by schedules of reinforcement. With multiple ratio schedules, research has shown an inverse relationship between ratio requirement and amount of tolerance that resulted from daily administration of the drug. In contrast, tolerance to the effects of cocaine on behavior under multiple interval schedules generally has developed regardless of interval value. Under interval schedules reinforcement depends on the animal making one response following a time interval. Thus, as time to respond increases, the time to reinforcement decreases. On the other hand, fixed ratio schedules require a specified number of responses to be made prior to reinforcement. Therefore, delaying the initiation of responding does not coincide with a significant decrease in the time to reinforcement. In the current experiment, 6 pigeons were trained to respond under a three-component multiple schedule, with a different tandem fixed-ratio 1 fixed-interval schedule in each component. The multiple schedule required one response, which was followed by one of three fixed-interval values (5, 15, or 60 s). Thus, the multiple schedule was interval-like because after the fixed-ratio 1, only one more response was required for reinforcement, but it was also ratio-like because the length of the pause at the beginning of each interreinforcer interval affected the time until the next reinforcer. Acute administration of cocaine generally resulted in dose-dependent decreases in responding. Chronic (i.e., daily) administration of a rate-decreasing dose resulted in tolerance patterns similar to those usually obtained with multiple ratio schedules. That is, the magnitude of tolerance was related inversely to schedule size. These results suggest that delay to reinforcement from the initial response may play a role in the development of schedule-parameter-related tolerance.     

Conditioned reinforcement and discrimination in second-order schedules

Pigeons were exposed to multiple second-order schedules in which responding on the “main key” was reinforced according to either a variable-interval or fixed-interval schedule by production of a brief stimulus on the “brief-stimulus key”. A response was required to the brief stimulus during its fourth (final) presentation to produce food; responses to the earlier brief stimuli indicated the extent to which the final brief stimulus was discriminated from preceding ones. Main-key response rates were higher in early components of paired brief-stimulus schedules, in which each brief stimulus was the same as that paired with reinforcement, than in comparable unpaired brief-stimulus or tandem schedules. Poor discrimination occurred between paired brief stimuli (Experiment I). When chain stimuli on the main key induced a discrimination between the first two and second two brief stimuli, the response-rate enhancement in the paired brief-stimulus schedule persisted (Experiment II). Rate enhancement diminished when the initial link of the chain included the first three components (Experiment IV). Eliminating the contingency between responding and brief-stimulus production also diminished rate enhancement (Experiment III). The results show that the discriminative and conditioned reinforcing effects of food-paired brief stimuli may be selectively manipulated and suggest that the reinforcing effects are modulated by other reinforcers in the situation.     

Choice between sequences of fixed-ratio schedules: effects of ratio values and probability of food delivery.

Pigeons were exposed to schedules of food delivery that consisted of two sequential fixed ratios. When alternative sequences provided two food deliveries per 50 responses, the schedule with the shorter initial fixed-ratio value was consistently preferred. Progressively reducing from 1.0 to .25 the probability of food delivery following completion of the second fixed ratio of the sequence with the shorter initial fixed ratio did not reduce preference for this sequence. Moreover, the sequence with the shorter initial fixed ratio also was preferred when the probability of food delivery following completion of the initial ratio in that sequence was progressively reduced from 1.0 to .5, although preference shifted to the alternative when the probability was reduced to 0. These findings suggest that the length of the initial fixed ratio was a primary determinant of choice. Subsequent manipulations demonstrated, however, that when the initial fixed ratios of the two alternatives were equal, changes in the ratio value and probability of food delivery following completion of the second fixed ratio lawfully affected choice.