Delayed backward conditioning of place preference induced by wheel running in rats

Backward conditioning of place preference has been obtained in hungry rats when wheel running (the rewarding US) is followed immediately by exposure to a distinctive chamber (the CS). We tested whether such backward conditioning would occur with a 10-min delay. In each of four paired exposures, a period of wheel running (2 or 22 h, in separate experiments) was followed by 10 min in the home cage and then 30 min in the CS chamber. Control rats were put in the CS chamber without wheel running. In other similar experiments, a 0-, 10-, or 30-min delay separated wheel running from exposure to the CS chamber. Reliable conditioned place preference (CPP) occurred when the delay was 0 or 10 but not 30 min. The strength of CPP decreased as the delay increased. These findings imply that the reward process initiated by wheel running remains active for some time after running stops. They support the view that the suppression of feeding produced by wheel running in hungry rats (activity anorexia) is mediated by this reward process.     

Simple and conditional visual discrimination with wheel running as reinforcement in rats.

Three experiments explored whether access to wheel running is sufficient as reinforcement to establish and maintain simple and conditional visual discriminations in nondeprived rats. In Experiment 1, 2 rats learned to press a lit key to produce access to running; responding was virtually absent when the key was dark, but latencies to respond were longer than for customary food and water reinforcers. Increases in the intertrial interval did not improve the discrimination performance. In Experiment 2, 3 rats acquired a go-left/go-right discrimination with a trial-initiating response and reached an accuracy that exceeded 80%; when two keys showed a steady light, pressing the left key produced access to running whereas pressing the right key produced access to running when both keys showed blinking light. Latencies to respond to the lights shortened when the trial-initiation response was introduced and became much shorter than in Experiment 1. In Experiment 3, 1 rat acquired a conditional discrimination task (matching to sample) with steady versus blinking lights at an accuracy exceeding 80%. A trial-initiation response allowed self-paced trials as in Experiment 2. When the rat was exposed to the task for 19 successive 24-hr periods with access to food and water, the discrimination performance settled in a typical circadian pattern and peak accuracy exceeded 90%. When the trial-initiation response was under extinction, without access to running, the circadian activity pattern determined the time of spontaneous recovery. The experiments demonstrate that wheel-running reinforcement can be used to establish and maintain simple and conditional visual discriminations in nondeprived rats.     

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.     

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 pre-operant running and sucrose concentration on operant wheel running on a fixed interval schedule of reinforcement

Prior research proposed that temporal control over the pattern of operant wheel running on a fixed interval (FI) schedule of sucrose reinforcement is a function of automatic reinforcement generated by wheel running and the experimentally arranged sucrose reinforcement. Two experiments were conducted to assess this prediction. In the first experiment, rats ran for different durations (0, 30, 60, and 180 min) prior to a session of operant wheel running on a FI 120-s schedule. In the second experiment, the concentration of sucrose reinforcement on a FI 180-s schedule was varied across values of 0, 5, 15, and 25%. In Experiment 1, as the duration of pre-operant running increased, the postreinforcement pause before initiation of running lengthened while wheel revolutions in the latter part of the FI interval increased. In Experiment 2, wheel revolutions markedly increased then decreased to a plateau early in the FI interval. Neither manipulation increased temporal control of the pattern of wheel running. Instead, results indicate that operant wheel running is regulated by automatic reinforcement generated by wheel activity and an adjunctive pattern of running induced by the temporal presentation of sucrose. Furthermore, the findings question whether the sucrose contingency regulates wheel running as a reinforcing consequence.     

Instrumental learning as a result of omission training on wheel running

The present experiment sought to provide unequivocal evidence of instrumental learning under omission training. Hungry rats received free food reinforcement while spontaneously running in a wheel. For an omission group, running postponed or cancelled reinforcers in the presence of a discriminative stimulus (SD) requiring subjects to reduce responding to earn food. Background food presentations were then yoked to reinforcement delivered in the presence of the discriminative stimulus. For a control group, which received the same stimulus presentations, reinforcement delivery was yoked to the experimental group at all times. The procedure allowed both within- and between-subject comparisons between omission and response-independent schedules. The response-reinforcer delay under the omission contingency was adjusted so as to equate reinforcement frequency in the presence and absence of the SD. As the SD was not correlated differentially with reinforcement and the running response did not involve approach or withdrawal to the site of food delivery, the successful discrimination performance observed in this experiment cannot be accounted for by appeal to implicit classical conditioning. Instead, it is suggested that decreased running in the presence of the discriminative stimulus was based on the animals' veridical representation of the negative contingency between the response and reinforcement.     

Taste avoidance caused by spontaneous wheel running: effects of duration and delay of wheel confinement

Confinement of a rat in a running wheel results in the rat's subsequent avoidance of the taste consumed before the confinement. This phenomenon has been ascribed to taste aversion conditioned by spontaneous wheel running. As a first step toward clarification of the underlying mechanism of this phenomenon, we manipulated two parameters of the taste-confinement procedure: duration of wheel confinement (Experiments 1A and 1B) and temporal intervals between the taste consumption and the wheel confinement (Experiments 2A and 2B). In general, longer confinement and shorter inter-event interval caused stronger taste avoidance. However, the results also suggested that it is possible to establish taste avoidance when wheel confinement was delayed 1-h after the taste consumption. These results correspond to those of conventional taste aversion caused by illness-inducing agents, suggesting similar mechanisms in the both preparations. Experiment 3 revealed that running in a wheel rather than wheel confinement itself is the effective factor for establishing taste avoidance.     

Scheduled running wheel activity indexes the specificity of pharmacological anorexia.

Nondeprived male Sprague-Dawley rats that were given scheduled access to running wheels for 60 min daily ran immediately and energetically. Intraperitoneal injections of 400 micrograms/kg pancreatic glucagon and 0.15 microgram/kg cholecystokinin octapeptide had no effect on scheduled running, but significantly inhibited feeding when the rats were offered condensed milk instead of access to the running wheels. This is consistent with the hypothesized function of these peptides as postprandial satiety signals. In contrast, 0.5 mg/kg amphetamine and 75 microM/kg LiCl, which produced similar degrees of anorexia, inhibited running by about 50%. Amphetamine, but neither peptide, also inhibited water drinking and disrupted the behavioral sequence of postprandial satiety. The distance run during scheduled running tests was inversely related to body weight, but the patterns of the drugs' effects were not altered by baseline running differences. Scheduled wheel running is a robust consummatory behavior that appears to provide a relatively valid, simple, and sensitive test of the behavioral specificity of pharmacological anorexia.     

Effects of voluntary wheel running on heart rate, body temperature, and locomotor activity in response to acute and repeated stressor exposures in rats

Stress often negatively impacts physical and mental health but it has been suggested that voluntary physical activity may benefit health by reducing some of the effects of stress. The present experiments tested whether voluntary exercise can reduce heart rate, core body temperature and locomotor activity responses to acute (novelty or loud noise) or repeated stress (loud noise). After 6 weeks of running-wheel access, rats exposed to a novel environment had reduced heart rate, core body temperature, and locomotor activity responses compared to rats housed under sedentary conditions. In contrast, none of these measures were different between exercised and sedentary rats following acute 30-min noise exposures, at either 85 or 98 dB. Following 10 weeks of running-wheel access, both groups displayed significant habituation of all these responses to 10 consecutive daily 30-min presentations of 98 dB noise stress. However, the extent of habituation of all three responses was significantly enhanced in exercised compared to sedentary animals on the last exposure to noise. These results suggest that in physically active animals, under some conditions, acute responses to stress exposure may be reduced, and response habituation to repeated stress may be enhanced, which ultimately may reduce the negative and cumulative impact of stress.     

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.     

Increased eating in rats deprived of running

Daily food intake in rats was temporarily reduced by the introduction of an activity wheel and temporarily increased by the subsequent removal of the wheel. When this outcome is coupled with the positive relation between food deprivation and running—and food deprivation is seen as a loss of eating rather than as a physiological state—there is the suggestion that the total behavior output of the organism may be regulated as such. Specifically, when the rat is deprived of a behavior that recurrently comprises a large part of its total daily activity, an increase may occur in some other behavior.     

Effects of glucose-saccharin preloads on schedule- and deprivation-induced drinking in rats

The present series of experiments found that high volumes of a glucose-saccharin solution consumed by rats prior to testing (i.e., a preload) produced a small, but significant suppression of established schedule-induced drinking (Experiment 2). This reduction in polydipsic drinking did not appear to be due to the caloric value of the preload (Experiment 1) or to any changes in food motivation, since Experiment 2 showed that there were no changes in lever pressing on a fixed-interval 1-min food schedule. In Experiment 3, the glucose-saccharin preloads produced a complete suppression of water deprivation-induced drinking. While schedule-induced drinking does appear to be a nonhomeostatic form of drinking, under certain testing conditions, it is sensitive to manipulations which affect homeostatic thirst mechanisms.