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.     

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.     

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).     

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.     

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.     

Flavor preferences produced by backward pairing with wheel running

In 3 experiments rats given 8 sessions of preexposure to wheel running acquired a preference for a flavor that was given immediately after each of 4 subsequent sessions of wheel running. Such flavor preference was less likely when rats were given the same conditioning procedure but without preexposure to wheels (Experiment 1) or when access to flavor was delayed by 30 min following a wheel session (Experiment 2). When rats were given a flavor before each wheel session, the resulting conditioned aversion was greater in rats that had no prior exposure to wheel running (Experiment 3). These results show that whether an aversion or preference for a flavor is produced by wheel running depends on an interaction between prior wheel experience and the sequence of events.     

A running wheel tachometer

A photocell-actuated tachometer for running wheels is described. The number of counts generated per revolution of the wheel is a function of the number of trip wires inserted in the hub of the wheel axle. The circuit is presented in configurations that actuate relay contact closures and as a direct driver of TTL logic.     

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.     

Some effects of punishment upon unpunished responding

Animals permitted free access to a running wheel and drinking tube increased the amount of running when drinking was punished with electric shock. Additional experiments demonstrated that the simple presence or absence of a drinking tube (or running wheel) was a sufficient condition to observe a decrease or an increase in the alternative response. A quantitative analysis of these interactions observed between the incompatible running and drinking responses suggested that each response occupied a constant proportion of the time available for it. These results question an interpretation of the increase in unpunished alternative responding based upon its avoidance properties.     

Gender dimorphic effects of voluntary running in laboratory rats depends on maturational status

This study examined the effect of 24 hr per day wheel access on running, body weight, and food intake for 30- or 50-day-old male and female rats under ad lib feeding conditions. Food intake and body weight were also monitored in a control group housed without access to running wheels. A dimorphic effect was observed after wheel introduction in 50-day-old but not 30-day-old rats: A temporary decline in food intake and a lasting decrease in body weight occurred for active male rats in comparison to their sedentary controls, and wheel access facilitated food intake and preserved body weight gain in female rats in comparison to their sedentary counterparts. Hyperphagia in adult females is interpreted in terms of the evolutionary acquired advantage linked to their reproductive function.     

Gender dimorphic effects of voluntary running in laboratory rats depends on maturational status

This study examined the effect of 24?hr per day wheel access on running, body weight, and food intake for 30- or 50-day-old male and female rats under ad lib feeding conditions. Food intake and body weight were also monitored in a control group housed without access to running wheels. A dimorphic effect was observed after wheel introduction in 50-day-old but not 30-day-old rats: A temporary decline in food intake and a lasting decrease in body weight occurred for active male rats in comparison to their sedentary controls, and wheel access facilitated food intake and preserved body weight gain in female rats in comparison to their sedentary counterparts. Hyperphagia in adult females is interpreted in terms of the evolutionary acquired advantage linked to their reproductive function.     

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.     

Flavour aversion produced by running and attenuated by prior exposure to wheels

In two experiments hungry rats were given access to running wheels. When given the novel flavour, almond, prior to novel access to the wheels, a conditioned aversion to almond was revealed by a subsequent two-bottle test. No such aversion was found in rats with previous experience of wheel running, whether this prior running occurred in the absence of any novel flavour, as in Experiment 1, or following access to saccharin, as in Experiment 2. These results suggest that the failure of rats with prior experience of the running wheels to develop a flavour aversion (unconditioned stimulus, US, preexposure effect) is unlikely to be due to associative blocking. Instead it seems that increasing exposure to a wheel produces habituation of its nausea-inducing properties.     

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.     

Reinforcement of wheel running in BALB/c mice: role of motor activity and endogenous opioids

The authors investigated the effect of the opioid antagonist naloxone on wheel-running behavior in Balb/c mice. Naloxone delayed the acquisition of wheel-running behavior, but did not reduce the expression of this behavior once acquired. Delayed acquisition was not likely a result of reduced locomotor activity, as naloxone-treated mice did not exhibit reduced wheel running after the behavior was acquired, and they performed normally on the rotarod test. However, naloxone-blocked conditioned place preference for a novel compartment paired previously with wheel running, suggesting that naloxone may delay wheel-running acquisition by blocking the rewarding or reinforcing effects of the behavior. These results suggest that the endogenous opioid system mediates the initial reinforcing effects of wheel running that are important in acquisition of the behavior.     

Intercurrent and reinforced behavior under multiple spaced-responding schedules

Lever pressing in rats was reinforced with food under a multiple spaced-responding schedule. A lever, food cup, and drinking tube were mounted in a running wheel so that lever pressing, running, and licking could be recorded. Running and licking had no scheduled consequences. Lever pressing was reinforced under a multiple schedule with three spaced-responding components and an extinction component. Each component was associated with a different auditory stimulus. Spaced-responding components reinforced only lever presses terminating interresponse times equal to or greater than 10, 20, or 60 sec, respectively. Rates of lever pressing, reinforcement, and licking all decreased as schedule parameter increased. Efficiency of spaced responding, as measured by reinforcements per response, also decreased. Rate of wheel running either increased or increased and then decreased with increasing schedule parameter. Individual running rates differed substantially. Neither licking nor running rate correlated with individual differences in efficiency. Analysis of conditional probabilities among the several response classes showed that, as the schedule requirement increased, the probability of running after a lever press increased and the probability of licking after a lever press decreased. After reinforcement, one subject always pressed the lever next. In the other subjects, the conditional probability of lever pressing, given reinforcement, increased while the probability of licking, given reinforcement, decreased with increasing schedule requirement. Results are discussed in relation to the concepts of schedule-induced and mediating behavior.     

Distributional properties of operant-level locomotion in the rat

Four rats had continuous access to activity wheels first, then access for 1 hr per day, and, subsequently, continuous access. Limiting S's access to the wheel substantially increased the total frequency of running. A distributional analysis of response duration, burst duration, and interburst interval showed that the increased frequency arose almost entirely from a shortening of the interval between successive bursts. In contrast, speed of the individual response and number of responses per burst changed only negligibly. If S were running, the probability that it would either stop or continue did not differ appreciably for the conditions of continuous or limited access to the wheel. But if S were not running, the probability that it would start running was appreciably greater for limited than for continuous access.     

Preadaptation to the feeding schedule does not eliminate activity-based anorexia in rats

To test whether activity-based anorexia (ABA) still occurs after preadaptation to the feeding schedule, 20 rats were first exposed to a feeding schedule of one 90-min meal per day until adaptation occurred (measured by maintenance of stable body weight). Then, during ABA training, half the rats (wheel group) were confined in running wheels except during the daily meal, and half (cage group) were not. Wheel running suppressed feeding—that is, food intake in the wheel group was less than that in the cage group. Also, the rats in the wheel group lost weight, whereas those in the cage group did not. Wheel running increased over days. Thus, the defining characteristics of ABA were evident in rats that were not subjected to ABA training until after they had become well adapted to the feeding schedule. These findings support the view that the suppression of feeding produced by wheel running triggers the vicious circle of ABA. They also cast doubt on the hypothesis that activity-induced interference with adaptation to the feeding schedule plays a key role in causing ABA.     

Activity patterns in rats (Rattus norvegicus) as a function of the cost of access to four resources

Patterns of eating, drinking, wheel running, and nesting were recorded in 2 experiments in which rats (Rattus norvegicus) lived in a laboratory environment that provided food, water, a running wheel, and a nest box. Access to each resource was contingent on the completion of a fixed ratio of bar presses and once earned remained available until the resource was not used for 10 consecutive min. In all cases an increase in the access price of a resource produced a decrease in the frequency with which the resource was accessed. This reduction in bout frequency was countered by an increase in bout size, which was compensatory for eating and nearly so for drinking, but which was only partially compensatory for wheel running. Nest bout size did not change significantly as nest price increased. The bout patterns of these 4 activities changed independently of one another, and the probabilities of behavioral transitions did not indicate strong links between any pairs of activities.     

A simple and sensitive method for monitoring running-wheel movement

A running-wheel movement-detection method is described for use with a modified 1350 Commodore mouse and a Commodore 64, is described. The movement-detection method allows for the detection of partial revolutions and direction of movement, but requires no interface equipment. The modified running wheel is discussed as a new technique that may be useful for bridging empirical and theoretical differences between free-operant and discrete-trial runway procedures.