Effects of post-session wheel running on within-session changes in operant responding

This study tested the effects of post-session wheel running on within-session changes in operant responding. Lever-pressing by six rats was reinforced by a food pellet under a continuous reinforcement (CRF) schedule in 30-min sessions. Two different flavored food pellets were used as reinforcers. In the wheel conditions, 30-min operant-sessions with one of the flavored pellets were followed by 30-min free-wheel running sessions. Meanwhile, in the home conditions, rats’ operant responding was reinforced by the other flavored pellets during 30-min operant-sessions, and the rats were then returned to their homecages. All rats were exposed to 4 wheel and 4 homecage sessions. Operant responding was lowered during the wheel conditions. However, post-session running did not alter the within-session pattern of operant responding. These effects were practically identical to the effects of drug-induced taste-aversion learning on within-session changes in operant responding, suggesting similar mechanisms in both taste-aversion preparations.     

Effects Of Fixed-interval Schedule And Reinforcer Duration On Responding Reinforced By The Opportunity To Run

Two experiments investigated the effects of schedule value and reinforcer duration on responding for the opportunity to run on fixed-interval (FI) schedules in rats. In the first experiment, 8 male Wistar rats were exposed to FI 15-s, 30-s, and 60-s schedules of wheel-running reinforcement. The operant was lever pressing, and the consequence was the opportunity to run for 60 s. In the second experiment, 8 male Long-Evans rats were exposed to reinforcer durations of 15 s, 30 s, and 90 s. The schedule of reinforcement was an FI 60-s schedule. Results showed that postreinforcement pause and wheel-running rates varied systematically with reinforcer duration but not schedule value. Local lever-pressing rates decreased with reinforcer duration. Overall lever-pressing rates decreased with reinforcer duration but increased with schedule value. Although the reinforcer-duration effect is consistent with previous research, the lack a schedule effect appears to be the result of long post-reinforcement pauses following wheel-running reinforcement that render the manipulation of the interval requirement ineffective.     

Deprivation and satiation: The interrelations between food and wheel running

Two experiments were designed to assess whether depriving rats of food would increase the reinforcement effectiveness of wheel running (Experiment 1) and whether satiation for wheel running would decrease the reinforcement effectiveness of food (Experiment 2). In Experiment 1, a progressive-ratio schedule was used to measure the reinforcement effectiveness of wheel running when rats were deprived or not deprived of food. Completion of a fixed number of lever presses released a brake on a running wheel for 60 s, and the response requirement was systematically increased until the rat stopped pressing or until 8 hr had elapsed. The ratio value reached (and the total number of lever presses) was an inverted-U function of food deprivation (percentage body weight). In Experiment 2, when wheel running preceded test sessions, fewer food-reinforced lever presses were maintained by the progressive-ratio schedule, and responding occurred at a lower rate on a variable-interval schedule. An interpretation of these results is that deprivation or satiation with respect to one event (such as food) alters the reinforcement effectiveness of a different event (such as access to wheel running).     

Reinforcement value and substitutability of sucrose and wheel running: implications for activity anorexia

Choice between sucrose and wheel-running reinforcement was assessed in two experiments. In the first experiment, ten male Wistar rats were exposed to concurrent VI 30 s VI 30 s schedules of wheel-running and sucrose reinforcement. Sucrose concentration varied across concentrations of 2.5, 7.5, and 12.5%. As concentration increased, more behavior was allocated to sucrose and more reinforcements were obtained from that alternative. Allocation of behavior to wheel running decreased, but obtained wheel-running reinforcement did not change. Overall, the results suggested that food-deprived rats were sensitive to qualitative changes in food supply (sucrose concentration) while continuing to defend a level of physical activity (wheel running). In the second study, 15 female Long Evans rats were exposed to concurrent variable ratio schedules of sucrose and wheel-running, wheel-running and wheel-running, and sucrose and sucrose reinforcement. For each pair of reinforcers, substitutability was assessed by the effect of income-compensated price changes on consumption of the two reinforcers. Results showed that, as expected, sucrose substituted for sucrose and wheel running substituted for wheel running. Wheel running, however, did not substitute for sucrose; but sucrose partially substituted for wheel running. We address the implications of the interrelationships of sucrose and wheel running for an understanding of activity anorexia.     

Effects of cocaine on fixed-interval responding reinforced by the opportunity to run

Rate-dependent drug effects have been observed for operant responding maintained by food, water, heat, light onset, electrical brain stimulation, shock-stimulus termination, and shock presentation. The present study sought to determine if the effects of cocaine on lever pressing maintained by the opportunity to run could also be described as rate dependent. Seven male Wistar rats were trained to respond on levers for the opportunity to run in a wheel. The schedule of reinforcement was fixed-interval 60 s, and the reinforcing consequence was the opportunity to run for 60 s. On this schedule, overall rates of responding were low, usually below six presses per minute, and pauses frequently exceeded the 60-s interval. Despite these differences, an overall scalloped pattern of lever pressing was evident for each rat. Doses of 1, 2, 4, 8, and 16 mg/kg cocaine were administered 10 min prior to a session. Only at the 16 mg/kg dose did the responding of the majority of rats change in a manner suggestive of a rate-dependent drug effect. Specifically, lower response rates at the beginning of the intervals increased and higher rates at the end of the intervals decreased, as indicated by the fact that slopes from the regression of drug rates on control rates decreased. These data provide tentative support for the generalization of rate-dependent effects to operant responding maintained by wheel running. Differences in the baseline performance maintained by wheel running compared to those for food and water point to the need for further experimentation before this effect can be firmly established.     

The Probability of Small Schedule Values and Preference for Random-Interval Schedules

Preference for working on variable schedules and temporal discrimination were simultaneously examined in two experiments using a discrete-trial, concurrent-chains arrangement with fixed interval (FI) and random interval (RI) terminal links. The random schedule was generated by first sampling a probability distribution after the programmed delay to reinforcement on the FI schedule had elapsed, and thus the RI never produced a component schedule value shorter than the FI and maintained a rate of reinforcement half that of the FI. Despite these features, the FI was not strongly preferred. The probability of obtaining the smallest programmed delay to reinforcement on the RI schedule was manipulated in Experiment 1, and the interaction of this probability and initial link length was examined in Experiment 2. As the probability of obtaining small values in the RI increased, preference for the schedule increased while the discriminated time of reinforcer availability in the terminal link decreased. Both of these effects were attenuated by lengthening the initial links. The results support the view that in addition to the delay to reinforcement, the probability of obtaining a short delay is an important choice-affecting variable that likely contributes to the robust preferences for variable, as opposed to fixed, schedules of reinforcement.     

Competitive fixed-interval performance in humans

Two persons responded in the same session in separate cubicles, but under a single schedule of reinforcement. Each time reinforcement was programmed, only the first response to occur, that is, the response of only one of the subjects, was reinforced. “Competitive” behavior that developed under these conditions was examined in three experiments. In Experiment 1 subjects responded under fixed-interval (FI) 30-s, 60-s, and 90-s schedules of reinforcement. Under the competition condition, relative to baseline conditions, the response rates were higher and the pattern was “break-and-run.” In Experiment 2, subjects were exposed first to a conventional FI schedule and then to an FI competition schedule. Next, they were trained to respond under either a differential-reinforcement-of-low-rate (DRL) or fixed-ratio (FR) schedule, and finally, the initial FI competition condition was reinstated. In this second exposure to the FI competition procedure, DRL subjects responded at lower rates than were emitted during the initial exposure to that condition and FR subjects responded at higher rates. For all subjects, however, responding gradually returned to the break-and-run pattern that had occurred during the first FI competition condition. Experiment 3 assessed potential variables contributing to the effects of the competitive FI contingencies during Experiments 1 and 2. Subjects were exposed to FI schedules where (a) probability of reinforcement at completion of each fixed interval was varied, or (b) a limited hold was in effect for reinforcement. Only under the limited hold was responding similar to that observed in previous experiments.     

Quantitative studies of reinforcement relativity

Three experiments examined an application of the matching law to the area of reinforcement relativity. In Experiment I, rats ran in a wheel and drank a sucrose solution. Equations derived from the matching law made fairly accurate predictions of the amounts of time spent running and drinking when licks and wheel revolutions had to occur in fixed proportions. In Experiment II, rats were required to spend four times as much time drinking as running, but the absolute durations of the cycles of drinking and running were varied. Except for the shortest cycle size tested, durations were close to those predicted. Experiments III investigated a tendency for obtained durations of running and drinking to be slightly longer than predicted. Simply shortening the periods when these behaviors were available increased their values. It was concluded that the matching law equations provided reasonably accurate predictions in some experiments, but changes in motivation set the limits of such accuracy.     

Exclusive preference develops less readily on concurrent ratio schedules with wheel-running than with sucrose reinforcement

Previous research suggested that allocation of responses on concurrent schedules of wheel‐running reinforcement was less sensitive to schedule differences than typically observed with more conventional reinforcers. To assess this possibility, 16 female Long Evans rats were exposed to concurrent FR FR schedules of reinforcement and the schedule value on one alternative was systematically increased. In one condition, the reinforcer on both alternatives was .1 ml of 7.5% sucrose solution; in the other, it was a 30‐s opportunity to run in a wheel. Results showed that the average ratio at which greater than 90% of responses were allocated to the unchanged alternative was higher with wheel‐running reinforcement. As the ratio requirement was initially increased, responding strongly shifted toward the unchanged alternative with sucrose, but not with wheel running. Instead, responding initially increased on both alternatives, then subsequently shifted toward the unchanged alternative. Furthermore, changeover responses as a percentage of total responses decreased with sucrose, but not wheel‐running reinforcement. Finally, for some animals, responding on the increasing ratio alternative decreased as the ratio requirement increased, but then stopped and did not decline with further increments. The implications of these results for theories of choice are discussed.     

Discriminative and reinforcing properties of two types of food pellets

In Experiment I some discriminative functions of food pellets were studied by developing a multiple schedule of reinforcement (mult FR 30 FI 3) in which the delivery of a standard laboratory food pellet as a reinforcer set the occasion for reinforcement on every 30th response (FR 30), and the delivery of a sucrose food pellet as a reinforcer set the occasion for reinforcement after a 3-min interval (FI 3). Discriminative stimulus control by the type of pellet was also demonstrated by reversing the operant discrimination and having the standard pellet control the FI 3 and the sucrose pellet control the FR 30. In Experiment II a mult FR 30 FR 30 with two bars was developed; a standard food pellet was followed by an FR 30 on Bar 1 and extinction (ext) on Bar 2, while a sucrose pellet was followed by an FR 30 on Bar 2 and ext on Bar 1. A control rat was placed, for comparison, on a mixed (mix) FR 30 FR 30 schedule with two bars, but neither bar correlated with the type of food pellet. In Experiments I and II the similarity between pellet controlled multiple schedules and multiple primed schedules was discussed, as was the comparability of transitions and effectiveness of control between pellet controlled multiple schedules and multiple schedules providing continuous exteroceptive stimuli.     

Habituation contributes to within-session changes in free wheel running.

Three experiments tested the hypothesis that habituation contributes to the regulation of wheel running. Rats ran in a wheel for 30-min sessions. Experiment 1 demonstrated spontaneous recovery. Rats ran more and the within-session decreases in running were smaller after 2 days of wheel deprivation than after 1 day. Experiment 2 demonstrated dishabituation. Running rate increased immediately after the termination of a brief extra event (application of the brake or flashing of the houselight). Experiment 3 demonstrated stimulus specificity. Rats completed the second half of the session in either the same wheel as the first half, or a different wheel. Second-half running was faster in the latter case. Within-session patterns of running were well described by equations that describe data from the habituation, motivation, and operant literatures. These results suggest that habituation contributes to the regulation of running. In fact, habituation provides a better explanation for the termination of wheel running than fatigue, the variable to which this termination is usually attributed. Overall, the present findings are consistent with the proposition that habituation and sensitization contribute to the regulation of several forms of motivated behavior.     

Techniques for establishing schedules with wheel running as reinforcement in rats.

In three experiments, access to wheel running was contingent on lever pressing. In each experiment, the duration of access to running was reduced gradually to 4, 5, or 6 s, and the schedule parameters were expanded gradually. The sessions lasted 2 hr. In Experiment 1, a fixed-ratio 20 schedule controlled a typical break-and-run pattern of lever pressing that was maintained throughout the session for 3 rats. In Experiment 2, a fixed-interval schedule of 6 min maintained lever pressing throughout the session for 3 rats, and for 1 rat, the rate of lever pressing was positively accelerated between reinforcements. In Experiment 3, a variable-ratio schedule of 20 or 35 was in effect and maintained lever pressing at a very stable pace throughout the session for 2 of 3 rats; for 1 rat, lever pressing was maintained at an irregular rate. When the session duration was extended to successive 24-hr periods, with food and water accessible in Experiment 3, lever pressing settled into a periodic pattern occurring at a high rate at approximately the same time each day. In each experiment, the rats that developed the highest local rates of running during wheel access also maintained the most stable and highest rates of lever pressing.     

Schedule constraint on the average drink burst and the regulation of wheel running and drinking in rats

Two experiments compared predictions of a molar-pattern model and a general molar behavior regulation model by requiring rats to wheel run for access to water. In both experiments schedule parameters constrained the baseline average burst length of drinking without constraining total drinking. Five levels of schedule constraint were imposed on time spent per drinking burst (Experiment 1) or the number of drinks per burst (Experiment 2). The results of both experiments supported the general molar behavior regulation view but not the molar-pattern model by showing no increase in total wheel running and no decrease in total drinking under schedule constraint. However, both experiments also showed local effects of drink burst constraint, including a direct relation between the degree of constraint and the local rate of drinking, and an approximation of the temporal distribution of baseline drinking under all degrees of schedule constraint. Most local changes support the view that rats defend the baseline temporal distribution of responding under schedule constraint, though some changes appear related to disruption of local response pattern characteristics.     

Multiple determinants of the effects of reinforcement magnitude on free-operant response rates

Four experiments examined the effects of increasing the number of food pellets given to hungry rats for a lever-press response. On a simple variable-interval 60-s schedule, increased number of pellets depressed response rates (Experiment 1). In Experiment 2, the decrease in response rate as a function of increased reinforcement magnitude was demonstrated on a variable-interval 30-s schedule, but enhanced rates of response were obtained with the same increase in reinforcement magnitude on a variable-ratio 30 schedule. In Experiment 3, higher rates of responding were maintained by the component of a concurrent variable-interval 60-s variable-interval 60-s schedule associated with a higher reinforcement magnitude. In Experiment 4, higher rates of response were produced in the component of a multiple variable-interval 60-s variable-interval 60-s schedule associated with the higher reinforcement magnitude. It is suggested that on simple schedules greater reinforcer magnitudes shape the reinforced pattern of responding more effectively than do smaller reinforcement magnitudes. This effect is, however, overridden by another process, such a contrast, when two magnitudes are presented within a single session on two-component schedules.     

Acquisition of a spatially defined operant with delayed reinforcement.

Two experiments investigated the role of an immediate, response-produced auditory stimulus during acquisition, via delayed reinforcement, of a response selected to control for possible unprogrammed, operandum-related sources of response feedback. Experimentally naive rats were exposed to a delayed-food reinforcement condition, specifically a tandem fixed-ratio 1 differential-reinforcement-of-other-behavior 30-s schedule. The response was defined as breaking a photocell beam located near the ceiling at the rear of the operant conditioning chamber. In Experiment 1, rates of photobeam breaking by each rat increased from near zero, regardless of the presence or absence of a tone that immediately followed the response initiating the delay interval. Though not essential, the tone facilitated response acquisition and resulted in more efficient response patterns at stability. Experiment 2 demonstrated that photobeam-breaking response rates under the delayed reinforcement contingency exceeded those in a preceding baseline condition in which no food was delivered. In addition, upon introduction of the delayed reinforcement procedure, correspondence between response patterns and the requirements of the reinforcement schedule increased over baseline levels in the absence of a food contingency. Together with a previous report of Lattal and Gleeson (1990), the present results suggest that response acquisition with delayed reinforcement is a robust phenomenon that may not depend on a mechanically defined response or an immediate external stimulus change to mediate the temporal gap between response and reinforcer.     

Effects of concurrent response-independent reinforcement on fixed-interval schedule performance

In three experiments, behavior maintained by fixed-interval schedules changed when response-independent reinforcement was delivered concurrently according to fixed- or variable-time schedules. In Experiment I, a pattern of positively accelerated responding during fixed interval was changed to a linear pattern when response-independent reinforcement occurred under a variable-time schedule. Overall response rates (total responses/total time) decreased as the frequency of response-independent reinforcement increased. Experiment II showed that the response-rate changes in the first experiment were controlled by the response-reinforcer relation, but the changes in patterns of responding were similar whether concurrently available reinforcement at varying times was response-dependent or response-independent. In the final experiment, the addition of response-independent reinforcement at fixed times to a fixed-interval schedule resulted in changes in both local and overall response rates and in the occurrence of positively accelerated responding between reinforcements. These results suggest that the temporal distribution of reinforcers determines response patterns and that both the response-reinforcement dependency and the schedule of reinforcement determine overall response rates during concurrently scheduled response-dependent and response-independent reinforcement.     

Diversity and substitutability of adjunctive activities under fixed-interval schedules of food reinforcement

Six rats received food contingent on pressing a lever on fixed-ratio 1, fixed-interval 30-second, and fixed-interval 60-second schedules, with concurrent access to a drinking spout, a running wheel, and a block of wood. Drinking, running, and chewing were monitored automatically, and these and other activities were observed directly during selected sessions. Because all sessions ended after delivery of 60 pellets, total time available for activities other than eating increased over the three schedules. Time spent contacting the lever and visiting the food tray increased in proportion to total available time, whereas the time spent in other activities changed in a complex manner such that drinking was the dominant adjunctive behavior in the 30-second condition, and running or chewing the dominant adjunctive behavior in five of six rats in the 60-second condition. General activity and grooming also occupied significant amounts of time. In a subsequent part of the experiment, running and chewing were prevented, and the majority of other activities, especially drinking and grooming, increased. The results show that (a) FI schedules of food reinforcement are accompanied by a wide variety of adjunctive activities; (b) the preferred activity differs according to the schedule duration; and (c) the extent to which activities substitute for one another is limited by the tendency for different activities to occupy different parts of the interreinforcement interval.     

Running and responding reinforced by the opportunity to run: effect of reinforcer duration.

The present study investigated the effect of reinforcer duration on running and on responding reinforced by the opportunity to run. Eleven male Wistar rats responded on levers for the opportunity to run in a running wheel. Opportunities to run were programmed to occur on a tandem fixed-ratio 1 variable-interval 30-s reinforcement schedule. Reinforcer duration varied across conditions from 30 to 120 s. As reinforcer duration increased, the rates of running and lever pressing declined, and latency to lever press increased. The increase in latency to respond was consistent with findings that unconditioned inhibitory aftereffects of reinforcement increase with reinforcer magnitude. The decrease in local lever-pressing rates, however, was inconsistent with the view that response strength increases with the duration of the reinforcer. Response rate varied inversely, not directly, with reinforcer duration. Furthermore, within-session data challenge satiation, fatigue, and response deprivation as determinants of the observed changes in running and responding. In sum, the results point to the need for further research with nonappetitive forms of reinforcement.     

Varying reinforcement magnitude on interval schedules

In Experiment 1, rats were trained on either a random-interval or a variable-interval 60-sec schedule of reinforcement, and reinforcement magnitude was varied across conditions between one and four pellets. Although the two schedules maintained different patterns of behaviour, patterns and rates of responding were not systematically affected by the variation in reinforcement magnitude. In Experiment 2, a regulated probability interval schedule that generated similar rates of reinforcement to those of the schedules of Experiment 1 was used, with the pattern of behaviour generated resembling that typical of a random-interval schedule. Changing reinforcement magnitude again produced few systematic changes in behaviour. In Experiment 3, a variable-ratio schedule was used within a procedure that otherwise resembled that of Experiments 1 and 2. Increasing the reinforcement magnitude now decreased the rates of responding, and examination of the patterns of responding showed that this came about because rates of responding were higher early in the interreinforcer interval in the one-pellet condition. These experiments demonstrate the insensitivity of behaviour under interval schedules to changes in reinforcement magnitude and suggest the operation of mechanisms different from those engaged by ratio schedules and discretetrial learning procedures.     

Effect of signaled reinforcement on response variability

Three experiments examined the effect of signaling reinforcement on rats' lever pressing on contingencies that reinforced variable responding to extend the exploration of signaled reinforcement to a schedule that has previously not been examined in this respect. In Experiment 1, rats responding on a lag-8 variability schedule with signaled reinforcement displayed greater levels of variability (U values) than rats on the same schedule lacking a reinforcement signal. In Experiment 2, rats responding on a differential reinforcement of least frequent responses schedule also displayed greater operant variability with a signal for reinforcement compared with rats without a reinforcement signal. In Experiment 3, a reinforcement signal decreased the variability of a response sequence when there was no variability requirement. These results offer empirical corroboration that operant variability responds to manipulations in the same manner as do other forms of operant response and that a reinforcement signal facilitates the emission of the required operant.