Weight Loss in Rats Produced by Running: Effects of Prior Experience and Individual Housing

To investigate factors affecting activity-based anorexia (ABA) or activity-stress (AS), rats were given 2-hr access to a running wheel immediately prior to their daily 1.5-hr food access during the light cycle. This produced a reduction in food intake, a steady increase in running, and a large drop in body weight with a prolonged delay before weight recovery began. Experiment 1 found that these effects were reduced in rats with prior experience of eating at this time of day. In contrast, prior experience of running in the wheel when on ad lib food enhanced these effects in Experiment 2, where a subsequent change for half the subjects to individual housing produced a further decrease in body weight. The latter factor was investigated from the outset of Experiment 3 and again individually housed rats showed greater weight loss than did group-housed rats. This experiment also found that in rats of the same age a low initial body weight predicts greater vulnerability to ABA. It was concluded that ABA results from activity-induced reduction of feeding, which prolongs adaptation to a new feeding schedule and is accentuated by social isolation.     

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).     

Effects of an activity-based anorexia procedure on within-session changes in nose-poke responding

This study tested the effects of an activity-based anorexia (ABA) procedure on within-session changes in responding. In the ABA group (N = 8), rats were given a 60-min feeding session and allowed to run in a running wheel for the remainder of each day. During the daily 60-min feeding session, each nose-poke response was reinforced by a food pellet. In the control group (N = 8), rats experienced the same procedure except that the wheel was locked and thus rats could not run. The experiment lasted for 6 days. Rats in the ABA group consumed less and lost more body weight than those in the control group. Within-session decreases in nose-poke responding were steeper for ABA than control rats. In addition, response rates were well described as linear functions of the cumulative number of reinforcers in both groups (r²s > .96). The regression lines for the ABA group had steeper slopes and smaller x-axis intercepts than those for the control group. However, the y-axis intercepts of the regression lines were similar for the two groups. These effects were different from the effects of taste-aversion learning induced by post-session wheel running (Aoyama, 2007), suggesting a different mechanism in each preparation.     

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.     

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.     

High ambient temperature reduces rate of body-weight loss produced by wheel running

This study examined the effect of ambient temperature (AT) on the relationship between activity and weight loss. Compared with a neutral AT of 21 degrees C, high ATs of 27-29 degrees C produced a slower rate of weight loss in rats given 1.5-hr food access and 22.5-hr running-wheel access in a standard activity-based anorexia (ABA) procedure (Experiments 1 and 2). The high AT did not affect food intake or wheel running in Experiment 1, but did reduce running in Experiment 2. Switching from neutral to high AT had only a transient effect on weight loss when wheel access was maintained (Experiment 2) but resulted in less weight loss when wheel access was prevented (Experiment 3). Giving rats only 3 hr of wheel access each day at a neutral AT also produced substantial weight loss, but less if for the rest of each day they were maintained at a high AT (Experiment 4).     

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.     

Infantile handling and the facilitation of discrimination and reversal learning

One group of rats was removed from their home cage and received daily handling from Days 3-21, while the control group of litter mates remained in the cage and did not receive any treatment. On Day 22 all rats were weaned and they were housed in individual cages until they were 60 days old. After two pretraining days, subjects were given daily blocks of one free and three forced trials in the T-maze and were rewarded with food after making the correct response. Following 20 days of training and testing on black-white discrimination, subjects were given 10 days of reversal training with four daily trials. Results indicate that the handled animals showed faster running and a greater number of correct choices than the control rats during both the acquisition and reversal learning phases of the experiment.     

High ambient temperature reduces rate of body-weight loss produced by wheel running

This study examined the effect of ambient temperature (AT) on the relationship between activity and weight loss. Compared with a neutral AT of 21°C, high ATs of 27–29°C produced a slower rate of weight loss in rats given 1.5-hr food access and 22.5-hr running-wheel access in a standard activity-based anorexia (ABA) procedure (Experiments 1 and 2). The high AT did not affect food intake or wheel running in Experiment 1, but did reduce running in Experiment 2. Switching from neutral to high AT had only a transient effect on weight loss when wheel access was maintained (Experiment 2) but resulted in less weight loss when wheel access was prevented (Experiment 3). Giving rats only 3 hr of wheel access each day at a neutral AT also produced substantial weight loss, but less if for the rest of each day they were maintained at a high AT (Experiment 4).     

Insulin and glucagon as determinants of body weight set point and microregulation in rats

Seven adult male rats were observed for body weight and microregulation (feeding, drinking, and running patterns) after manipulation of insulin and glucagon levels. They received three injections per day for 3 days each week of 3 U of protamine zinc insulin, .25 mg of zinc glucagon, 50 microgram of protamine zinc somatostatin (SRIF), or protamine zinc vehicle. Diabetes was then induced with an iv injection of streptozotocin (65 mg/kg), and the injection schedule was repeated after the full diabetic syndrome emerged. In all rats whose insulin levels were increased relative to glucagon levels, body weight increased; in those whose glucagon levels were increased relative to insulin levels, body weight decreased. All injections except vehicle reduced meal sizes in both normal and diabetic rats, but only insulin increased the frequency of feeding. These effects could be predicted by the glucostatic theory of food intake regulation and are thus interpreted as supportive of this theory. These results also support the hypothesis that the relative concentration of insulin to glucagon is a regulator of body weight set point.     

Effect of auditory isolation on activity-based anorexia

The objective was to study male and female rats exposed to Activity-Based Anorexia (ABA; simultaneous exposition to food restriction and free access to an activity wheel) under two conditions of sound attenuation, by means of different arrangements of wheels (standard versus isolation) in the laboratory room. Regardless of the sound attenuation condition, all but one male and one female with access to wheels had to be removed from the experiment, but extended ABA endurance was found in rats in the sound attenuation condition. Furthermore, significantly lower levels of running were observed in both males and females under the sound attenuation arrangement in the isolation condition. The results suggest that external stimulation plays a role in the activity displayed by rats exposed to ABA, and that the reduction of external stimulation diminishes running but does not protect rats from developing ABA.     

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.     

Disrupted patterns of consummatory behavior in rats with fornix transections

The feeding and drinking behavior was examined in male rats with fornix transections and sham-operated control rats. Total food and water consumption was recorded but supplemented by a pattern analysis of feeding and drinking behavior. The behavior of the rats was continuously monitored during four hour morning and afternoon sessions under ad lib access and during a two hour session following adaptation to a restricted access feeding schedule. Rats with fornix transections were more active and exhibited increased frequencies of rearing, eating and drinking. The increased meal frequency in rats with fornix transections was accompanied by decreased meal durations and a reduction in the length of intermeal intervals. Total food and water consumption was unaffected by fornix transection as were the duration of sleep bouts and the frequencies of grooming, sleeping and carrying shavings. Fornix transections also reduced food carrying and food hoarding but only under conditions of restricted food access. The results suggest that fornix transection does not alter major homeostatic regulatory mechanisms nor does it alter the components of feeding and drinking behavior. Fornix transection alters, instead, the organization of microregulatory feeding and drinking patterns.     

The currency of procurement cost

As the number of instrtumental responses required to procure access to food is increased, animals decrease the frequency of initiating meals and increase meal size, conserving total intake while limiting the increase in the overall cost of feeding. In two studies, one using wheel turns and one using bar presses as the instrumental response, we asked whether freely feeding laboratory rats measure cost according to the energy or the time they expend. In each study we varied both the price (i.e., number of wheel turns or bar presses) and the force required to make a response (i.e., torque on the wheel or weight of the bar). Price affected both procurement time (from the first to the last procurement response) and procurement work, whereas torque and bar weight affected work without altering time in most cases. Meal patterns were altered by all manipulations of price, but changes in torque and bar weight had little effect on meal patterns, except in the conditions in which they altered procurement time. These results suggest that time is a critical currency of procurement cost in rats.     

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.     

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.     

The stability and control of conditioned noise aversion in the tilt cage

Twenty-six rats were used to study the development of stable conditioned responding to terminate white noise (100 or 105 db) in a tilt cage apparatus. Preliminary work, with one end of the cage consistently associated with silence, demonstrated stability at a high level of cumulative noise-off time per session. Time spent at the noise-off cage end was unaffected by lay-off and showed adaptation to reversal of the cage end functions. Variability in reversal acquisition, insensitivity to stimulus intensity changes and persistence in remaining at one end of the cage in the absence of noise-termination contingencies indicated the need for more precise behavioral control. A schedule programming several reversals of the no-noise end at variable intervals within each session was developed. Performance on the latter schedule was characterized by stability of high noise-off times and also proved more sensitive than the preliminary method to stimulus intensity changes (55, 70, 100 db) as measured by both cumulated noise-off time and cage crossing rates. The ease of training combined with sensitivity to stimulus change and the lack of highly specific motor response requirements make this technique promising for use with animals potentially debilitated by drugs or surgical after-effects.