Suppression of operant behavior and schedule-induced licking in rats

The first experiment studied the effects of punishment on rats' lever pressing maintained by a fixed-interval schedule of food reinforcement and on the associated schedule-induced licking. When licking was followed by shock, licking was suppressed but lever pressing was largely unaffected. When lever pressing was followed by shock, lever pressing was suppressed but licking was unaffected. In both cases, the punished behavior recovered its previous unpunished level when the shocks were discontinued. In a second experiment, the rats' lever pressing was maintained by a variable-interval schedule of food reinforcement under which polydipsic licking also developed. Both lever pressing and licking were partially suppressed during a stimulus correlated with occasional unavoidable electric shocks. With a higher shock intensity, both behaviors were suppressed further. Both lever pressing and licking recovered their previous levels when shocks were discontinued. These results show that schedule-induced licking, which has been described as adjunctive behavior, can be suppressed by procedures that suppress reinforced lever pressing, an operant behavior.     

Schedule-induced licking during multiple schedules

Schedule-induced polydipsia was studied in rats bar pressing under two-component multiple schedules of food reinforcement. The first component of the multiple schedule was a variable-interval 1-min schedule throughout the experiment. The schedule comprising the second component was varied over blocks of sessions in terms of rate and magnitude of reinforcement, and was either variable-interval 3-min (one pellet), variable-interval 3-min (three pellets), variable-interval 1-min (one pellet), or extinction. Water intake per session varied with the rate of reinforcement in the schedule comprising the second component and was highest when the schedule was variable-interval 1-min. Both bar-pressing behavior and licking behavior showed behavioral interactions between the two components of the multiple schedules. With magnitude of reinforcement held constant, a matching relationship was observed between lick rate and reinforcement rate; the relative frequency of licks in the constant component matched the relative frequency of reinforcement in that component. Bar pressing, however, showed only a moderate degree of relativity matching. During the schedule-induced licking, a burst of licking followed each delivery of a pellet (post-prandial drinking). The duration of these bursts of licking was observed to be a function of the inter-reinforcement interval.     

Stimulus generalization of schedule-induced polydipsia

Five rats were exposed to an intradimensional discrimination by associating two tones of different frequency with the components of a multiple random-time 30-sec, extinction schedule of food presentation. After schedule-induced polydipsia developed and the intermittent schedule of food presentation established stable differential licking rates during the stimuli associated with the multiple schedule, a stimulus generalization test was conducted. When generalization testing was conducted by presenting stimuli that varied on the frequency dimension during the random-time 30-sec component of the multiple schedule, all five rats demonstrated moderately sloping symmetrical gradients. Thus, schedule-induced polydipsia can be brought under the control of stimuli other than the food pellet.     

The opponent-process theory of motivation: VIII. Quantitative and qualitative manipulations of food both modulate adjunctive behavior

The present studies extend the opponent-process theory of acquired motivation to the schedule-induced polydipsia paradigm and more generally to adjunctive behavior. The theory suggests that manipulations of the quality and the size of the pellet presented on an intermittent schedule should be important modulators of polydipsia. In Experiment 1, presenting animals with preferred, less preferred, or least preferred food pellets on a fixed-time 120-sec schedule resulted in progressively lower levels of water intake. In Experiment 2, the quality and size of the pellet were manipulated factorially. These variables were found to interact in the control of both the development and maintenance of schedule-induced drinking. Thus, predictions from the opponent-process theory were supported by these findings.     

Effects of response-dependent and independent electric shock on schedule-induced polydipsia

In Experiment I, lever pressing by rats was maintained by the delivery of food pellets under a 45-sec fixed-interval schedule. Fixed-time 180-sec and fixed-interval 180-sec schedules of shock delivery were systematically superimposed on the baseline food schedule to study effects on schedule-induced water intake. Response-dependent shock had little, if any, effect on water intake, whereas shocks independent of lever pressing attenuated fluid intake. In Experiment 2, rats received food pellets under a fixed-time 60-sec schedule. Electric shock delivered concurrently under a variable-time 180-sec schedule, but never while the animal was licking or within 5 sec after licking terminated, led to similar attenuation of water intake. These findings suggest that schedule-induced polydipsia is sensitive to differences in the functional properties of response-independent and dependent electric shock.     

Control of schedule-induced polydipsi: type, size, and spacing of meals

Rats were given daily 1-min variable-interval sessions for several types of food delivered in various amounts per reinforcement and the concurrent, schedule-induced polydipsia was measured. Dry, solid food was neither a necessary nor sufficient condition for the development of polydipsia. Small portions of liquid Standard Monkey Diet produced polydipsia, but 45-mg dextrose or sucrose pellets did not. Within the range studied, smaller portions of both solid and liquid foods produced more drinking than larger portions per reinforcement. Two-min variable-interval sessions produced a greater polydipsic response than 1-min variable-interval, even though the number of 45-mg Noyes pellets allowed per session was held constant. Polydipsia was greatly attenuated on these schedules when the number of pellets remained constant, but were delivered two at a time. Within the ranges studied, the concurrent polydipsic response was increased by decreasing the rate of food acquisition, either by using smaller portions of food per reinforcement or by increasing the interreinforcement time.     

Context dependent changes in the reinforcing strength of schedule-induced drinking.

Previous experiments show that the opportunity to engage in schedule-induced responding is reinforcing. In this experiment, the reinforcing strength of schedule-induced drinking was measured. Four rats were trained on a concurrent-chain schedule. The two terminal links provided food pellets on identical fixed-time schedules. In addition, one terminal link also provided the opportunity to press a button that operated a water dipper. In this link, the rats showed polydipsic drinking. Button-pressing rate for polydipsic drinking was a bitonic function of pellet rate, and it was possible to describe the relationship with a slightly modified version of the matching equation for primary reinforcement. This equation also closely fit the data from other studies. Initial-link response rates, however, did not appear to be influenced by the availability of water in the terminal links. Control conditions suggested that the reinforcing strength of polydipsia was strongly bound to the context provided by periodic food reinforcement.     

Effects of diazepam on schedule-controlled and schedule-induced behavior under signaled and unsignaled shock.

Schedule-controlled lever pressing and schedule-induced licking were studied in rats under a multiple fixed-interval fixed-interval schedule of food reinforcement upon which was superimposed a multiple variable-time variable-time schedule of electric-shock delivery. Shocks were signaled in one component of the multiple schedule and unsignaled in the other. The effects of diazepam upon the suppression of behavior during the signal (conditioned suppression) and during signaled and unsignaled shock (differential suppression) were studied under several shock intensities (Experiment 1) and at increased body weight (Experiment 2). In each study, diazepam led to dose-dependent increases in the rate of pressing and licking during signaled and unsignaled shock, but had little effect on conditioned suppression. the rate-enhancing effects of diazepam depended upon the intensity of shock, nature of the response, and whether or not shocks were signaled. The data was discussed in terms of (1) implications for understanding the effects of signaled and unsignaled shock on behavior, (2) the effects of diazepam on behavior suppressed by response-independent shock, and (3) comparison between operant and schedule-induced behavior.     

Suppression of schedule-induced drinking and food-reinforced bar pressing by tail-pinch is not reversed by naloxone

In Experiment 1 eleven food-deprived rats were tested in a schedule-induced drinking paradigm under both tail-pinch and non-tail-pinch conditions. Tail-pinch produced a strong suppression of schedule-induced drinking, licking, licks per milliliter, bar presses, and number of reinforcers received during 30-min test sessions. Experiment 2 showed that the narcotic antagonist naloxone (2 and 4 mg/kg) did not reverse the tail-pinch suppression of schedule-induced drinking and food-reinforced bar pressing. Experiment 3 demonstrated that the suppression of schedule-induced drinking by tail-pinch could not be attributed to a suppression of drinking behavior in general, as tail-pinch had no effect on deprivation-induced drinking. The failure of naloxone to reverse this blockade suggests that endogenous opiate systems do not play a very important role in the suppression of schedule-induced polydipsia by tail-pinch. It was suggested that the combination of both the schedule-induction paradigm and the tail-pinch procedure increased arousal levels to such a high level that all behaviors were suppressed.     

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.     

The effects of delayed reinforcement on variability and repetition of response sequences

Four experiments examined the effects of delays to reinforcement on key peck sequences of pigeons maintained under multiple schedules of contingencies that produced variable or repetitive behavior. In Experiments 1, 2, and 4, in the repeat component only the sequence right-right-left-left earned food, and in the vary component four-response sequences different from the previous 10 earned food. Experiments 1 and 2 examined the effects of nonresetting and resetting delays to reinforcement, respectively. In Experiment 3, in the repeat component sequences had to be the same as one of the previous three, whereas in the vary component sequences had to be different from each of the previous three for food. Experiment 4 compared postreinforcer delays to prereinforcement delays. With immediate reinforcement sequences occurred at a similar rate in the two components, but were less variable in the repeat component. Delays to reinforcement decreased the rate of sequences similarly in both components, but affected variability differently. Variability increased in the repeat component, but was unaffected in the vary component. These effects occurred regardless of the manner in which the delay to reinforcement was programmed or the contingency used to generate repetitive behavior. Furthermore, the effects were unique to prereinforcement delays.     

Spaced food but not electrical brain stimulation induces polydipsia and air licking.

An attempt was made to induce polydipsia in rats whose lever pressing was reinforced with food pellets or electrical brain stimulation. Nine food-deprived, water-sated rats drank water excessively during sessions in which food pellets were delivered. When brain stimulation was substituted for food, drinking immediately ceased. Delivering brain stimulation according to a variety of schedules, pairing brain stimulation with food reinforcement, and substituting an air stream for water, each failed to produce polydipsic licking. These results show that polydipsia is not induced by all reinforcers.     

THE MOTIVATIONAL PROPERTIES OF SCHEDULE-INDUCED POLYDIPSIA1

Schedule-induced polydipsia occurred during initial magazine training to Noyes pellets (45 mg), disappeared when lever-pressing was acquired on a continuous reinforcement schedule (CRF), and reappeared when the food contingency was changed to a 1-min variable interval schedule (VI 1 min). Polydipsia also developed under a VI 1 min food schedule when water was concurrently available on various fixed ratios (FR), rather than being freely available. The level of the polydipsia and its motivating properties allow it to be classified as a form of adjunctive behavior.     

Effects of signaled and unsignaled shock on schedule-controlled lever pressing and schedule-induced licking: Shock intensity and body weight

Schedule-controlled lever pressing and schedule-induced licking were studied in rats under a multiple fixed-interval fixed-interval schedule of food reinforcement. Following acquisition of stable rates of pressing and licking, a multiple variable-time variable-time schedule of electric-shock delivery was superimposed upon the baseline schedule. In only one component of the multiple schedule, a 5-sec stimulus preceded each shock (signaled shock). In the other component shock was unsignaled. Several shock intensities (Experiment 1) and body weights (Experiment 2) were studied. Lever pressing and licking were affected similarly by experimental manipulations, although with parametric differences. Depending upon shock intensity and body weight, rates of lever pressing and licking were hardly suppressed, suppressed primarily in the unsignaled shock component (differential suppression), or markedly suppressed in both components. Differential suppression during components with signaled and unsignaled shock and conditioned suppression of responding during the preshock stimulus appeared not to be functionally related. Differential suppression depended more on the discriminability of shock-free time, and on shock intensity, body weight, and the type of response than on the “preparatory” behavior preceding shock.     

The effects of signalled and unsignalled lick-dependent delays on the development of schedule-induced drinking in rats

Food pellets were programmed to be delivered to rats every 60 sec (Fixed Time 60-sec schedule), and the development of schedule-induced drinking was measured in terms of the amount of water consumed and the number of licks per inter-pellet interval. For some rats (masters) 10-sec delays in food delivery were dependent on licks. Yoked-control rats received food at the same time as their masters and independently of their own behaviour. In Experiment 1, in which the delays were signalled by a blackout, the master rats began to drink, but this schedule-induced behaviour then decreased to levels lower than those shown by the yoked controls. When the signalled delays were discontinued, the drinking of the master rats recovered. In Experiment 2, in which the delays were not signalled, the master rats did not develop as much schedule-induced drinking as the yoked controls, and discontinuing the delays led to only small increases in drinking. These results support the view that schedule-induced drinking is subject to control by its consequences.     

Effects of electric-shock delivery on schedule-induced water intake: delay of shock, shock intensity, and body-weight loss.

In each of four experiments, schedule-induced water intake in the rat was studied under fixed-time 40-sec food delivery. Experiments I and II studied the temporal relationship between response-independent electric-shock delivery and licking. Shock was delivered under a variable-time 60-sec schedule. A lick-dependent delay was imposed so that licking and shock delivery were systematically separated in time by a minimum of 1 to 15 sec. Over a wide range of shock intensities the data failed to reveal a consistent delay-of-shock effect. Similar shock intensities led to similar reduction of water intake at each delay of shock interval. Experiments III and IV studied the effects of body-weight loss on water intake during independent shock delivery. In Experiment III, shock was delivered under variable-time 60-sec with a minimum separation between shock and licking of 5 sec. In Experiment IV, shock was delivered under variable-time 180-sec. The minimum separation between shock and licking was 10 sec. In each study, the resistance of water intake to suppression by shock delivery increased as the degree of body-weight loss increased. Schedule-induced water intake was affected more by shock when the animal was maintained at 90% of free-feeding weight than at 70%.     

Food-deprivation effects on punished schedule-induced drinking in rats.

Food-deprived rats (at 80% of their free-feeding weights) were exposed to a fixed-time 60-s schedule of food-pellet presentation and developed schedule-induced drinking. Lick-dependent signaled delays (10 s) to food presentation led to decreased drinking, which recovered when the signaled delays were discontinued. A major effect of this punishment contingency was to increase the proportion of interpellet intervals without any licks. The drinking of yoked control rats, which received food at the same times as those exposed to the signaled delay contingency (masters), was not consistently reduced. When food-deprivation level was changed to 90%, all master and yoked control rats showed decreases in punished or unpunished schedule-induced drinking. When the body weights were reduced to 70%, most master rats increased punished behavior to levels similar to those of unpunished drinking. This effect was not observed for yoked controls. Therefore, body-weight loss increased the resistance of schedule-induced drinking to reductions by punishment. Food-deprivation effects on punished schedule-induced drinking are similar to their effects on food-maintained lever pressing. This dependency of punishment on food-deprivation level supports the view that schedule-induced drinking can be modified by the same variables that affect operant behavior in general.     

Body Weight Reduction by Water Deprivation and Schedule-Induced Phenomena

On of the most important factors in the generation of schedule induced polydipsia, and schedule induced self-injection of drugs is the prior reduction of the animals' body weight to 80% of their free-feeding weight by food deprivation. It is suggested that reductio of body weight to 80% by water deprivation should produce similar results. The series of experiments reported here showed that rats reduced to 80% body weight by water deprivation did not develop either schedule-induced polydipsia or schedule induced wheelrunning. The results thus suggest that body weight reduction may not be a critical condition in the development of schedule-induced phenonmena.     

A comparison of operant licking and lever pressing in the rat

Two groups of rats were trained on a signaled, free-operant, avoidance procedure to lick or to lever press in order to avoid shock while water-deprived or satiated and, in the case of licking, while ingesting deionized water, isotonic saline, or 10% sucrose. The most effective avoidance licking occurred while the rats were water-deprived and ingesting 10% sucrose. Water deprivation level had no effect on lever pressing for shock avoidance. Two other groups of rats were operantly conditioned to lick or to lever press for food pellets while waterdeprived or satiated and, in the case of licking, while ingesting deionized water or 10% sucrose. The most effective licking for food reinforcement occurred while the rats were water-deprived and ingesting 10% sucrose. Water deprivation level had no effect on lever pressing for food reinforcement. The data indicated that effective operant licking must be supported by factors related to water regulation and taste palatability.