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.     

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.     

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.     

Punishment of schedule-induced drinking in rats by signaled and unsignaled delays in food presentation

Food-deprived rats were exposed to a fixed-time 60-s schedule of food-pellet presentation and developed schedule-induced drinking. Using an ABA reversal design, three experiments investigated the effects of events then made dependent on licks. In Experiment 1, lick-dependent signaled delays (10 s) in food presentation in general led to decreased drinking, which recovered when the signaled delays were discontinued. The drinking of yoked-control rats, which received food at the same times as those exposed to the signaled-delay contingency, showed much smaller changes. Experiment 2 showed that 10-s lick-dependent signals alone did not reduce drinking. In Experiment 3, when licks produced unsignaled 10-s delays in food there were less marked and more gradual changes in drinking than in Experiment 1, although these effects again were greater than with yoked-control animals. We concluded that both signaled and unsignaled delays functioned as punishers of drinking. These findings support the view that schedule-induced drinking, like operant behavior, is subject to control by its consequences.     

Effects of haloperidol on schedule-induced polydipsia.

In dose-related amounts, the drug haloperidol attenuated schedule-induced drinking by rats prefed with 0.01-mg drug added to 0, 25, 50, 75 or all of 100 Noyes 45-mg pellets. Drug pellets also induced less drinking than did regular Noyes pellets by rats that obtained these pellets at 1-min intervals by bar pressing. Haloperidol also reduced bar pressing and, temporarily, rate of reinforcement. The results appeared not to be due to a general sedative effect of haloperidol but to its selective power to reduce angiotensin-induced drinking. Thus, schedule-induced drinking, which is abnormal in not causing satiation, is controllable by a drug that interferes with the renin-angiotensin hormone system thought to regulate normal drinking.     

Schedule-induced polydipsia suppresses pituitary-adrenal activity in rats

The effects of schedule-induced polydipsia (SIP) on pituitary-adrenal activity, as indicated by plasma levels of corticosterone, were examined in a series of experiments. Male (Experiments 1 and 3) and female (Experiment 2) rats were reduced to 80% of their free-feeding weight and given daily sessions on an intermittent-feeding schedule (fixed time of 60 sec). Half of the subjects in each experiment had water available during experimental sessions and the other half did not. Animals with water available in the experimental chamber exhibited SIP in all three experiments. In Experiment 1, blood samples were collected following (a) food consumption in the home cage, (b) a session on FT 60 sec, and (c) a session with pellets available in a cup in the experimental chamber. In Experiment 2, blood samples were taken prior to and following an FT 60-sec session, and following a session with pellets available in a cup in the chamber. In Experiment 3, pre- and postsession samples were obtained as in Experiment 2 (Part A). Subsequently, the opportunity to drink during sessions was removed, and the effect on corticoids was examined (Part B). The results indicate that (a) schedule-induced drinking suppresses pituitary-adrenal activity, (b) corticoid suppression may become a conditioned response to drinking in the chamber, and (c) corticoids return to presession levels following removal of water from the chamber. In view of these findings, it is hypothesized that SIP may serve an arousal-reducing role in intermittent-feeding situations.     

Tail-pinch hyperalgesia and analgesia: Test-specific opioid and nonopioid actions

Reactivity to noxious stimuli in rats is altered following acute exposure to tail-pinch. However, while our laboratory has reported that tail-pinch produces hyperalgesia as measured by the flinch-jump test and attenuates analgesic responses following morphine and cold-water swims, others have found that tail-pinch elicits an opioid-sensitive analgesia on the hot plate test and a nonopioid-sensitive analgesia on the writhing test. The first experiment of the present study examined whether tail-pinch altered responses on two somatic pain tests and showed that tail-pinch significantly decreased both jump thresholds and tail-flick latencies. In assessing whether tail-pinch hyperalgesia on the jump test was mediated by endogenous opioids, the second experiment indicated that low (0.1 and 0.5 mg/kg) doses of naloxone eliminated tail-pinch hyperalgesia by selectively lowering control thresholds and a high (10 mg/kg) dose of naloxone eliminated tail-pinch hyperalgesia by selectively increasing thresholds following tail-pinch. Further, the third experiment showed that morphine-tolerant rats (10 mg/kg morphine daily over 14 days) did not exhibit tail-pinch hyperalgesia on the 15th day, an effect attributable to a selective lowering of control thresholds. The fourth experiment was a direct replication of the observation that tail-pinch produces analgesia on the writhing test which is not antagonized by naloxone. These results demonstrate that the pain test employed and the amount of prior tail-pinch experience are critical variables in determining the direction of tail-pinch effects upon pain perception in rats.     

Are schedule-induced drinking and displacement activities causally related?

The acquisition of schedule-induced drinking was studied in hungry rats given food according to an intermittent (fixed-time 45-s) schedule, in three separate experiments. Rate of acquisition of induced drinking was not affected by making the rats thirsty as well as hungry (Experiment I), nor by offering them a palatable saline solution instead of water (Experiment II). Thirst and palatability did affect the asymptotic level of induced drinking. Increasing the level of food deprivation (Experiment III) increased both the rate of acquisition of induced drinking and its asymptotic level. Thus, rate of acquisition of induced drinking was found to depend more on causal factors relevant to feeding than on causal factors relevant to drinking, while asymptotic level of drinking was affected by both types of causal factor. This suggests that the occurrence of induced drinking does not depend primarily on a disinhibition mechanism of the type thought to underlie displacement activities, and hence does not support the idea that induced drinking is a type of displacement activity.     

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.     

Stimulus- and pellet-induced drinking during a successive discrimination

In three experiments, interim water drinking was examined in rats exposed to a multiple schedule whose two components were extinction and a variable-time 30-s schedule of food delivery. Two different drinking patterns were observed in Experiment 1. Pellet-induced drinking, characterized by high rates of postpellet drinking in the variable-time component, with little or no drinking in extinction, occurred when the acquisition of stable postpellet drinking preceded discrimination training. Stimulus-induced drinking, characterized by a burst of drinking at the onset of extinction, with no drinking during the variable-time schedule, occurred when discrimination training preceded all other experimental conditions. With extended training, stimulus-induced drinking eventually was accompanied by postpellet drinking. In Experiment 2, the rate of stimulus-induced drinking and the number of sessions during which it occurred without postpellet drinking were found to be inversely related to component duration. In Experiment 3, the rate of schedule-induced drinking was found to vary directly with component duration.     

Naloxone and Pavlovian fear conditioning

Previous research has shown that pretreating rats with the opiate antagonist naloxone increases the freezing that follows painful electric shock. Three experiments, using freezing behavior as a dependent variable, were carried out to determine whether the drug might cause this effect by enhancing fear conditioning. Two of the studies employed a differential context fear-conditioning paradigm. Naloxone did not affect freezing behavior during the preshock adaptation period. In Experiment 1, naloxone was found to increase resistance to extinction in the S+ context. In Experiment 2, naloxone was found to increase freezing in the S+ context. This effect was dependent upon administering naloxone during training but not dependent on administering it during testing. The third study employed a generalization paradigm. It was found that naloxone's effect on postshock freezing was dependent on the place of testing; as the contextual cues of the test chamber were changed from those of the conditioning chamber, the effect of naloxone on freezing was reduced. The results of these experiments lend strong support to the hypothesis that naloxone increases freezing by enhancing the conditioning of fear to contextual stimuli associated with shock.     

Behaviour of Rats During Selfinitiated Pauses in Feeding and Drinking, and During Periodic Response-Independent Delivery of Food and Water

In Experiment I rats deprived of either food or water were given free access to food or water respectively, and their behaviour was observed during self-imposed pauses in feeding or drinking. In Experiment II food or water were delivered according to fixed-time 30-s and fixed-time 60-s schedules, and the behaviour of the rats was observed during the interreinforcement intervals imposed by these schedules. In both experiments the temporal pattern of those activities that occurred during pauses in eating differed from the pattern of activities occurring during pauses in drinking; and with both food and water the temporal pattern of activities during self-imposed pauses in consummatory behaviour in Experiment I proved a good predictor of the pattern of activities during schedule-imposed interreinforcement intervals in Experiment II. This suggests that intermittent schedules permit the occurrence of those activities that are normally closely associated with the consummatory behaviour in question. In Experiment II certain activities that occurred towards the end of the interreinforcement interval were found to be enhanced relative to baseline level, but there was no enhancement of activities occurring near the beginning of the interval. This is contrary to Staddon's (1977) account of schedule-induced behaviour, and suggests that schedule-induction is not as common as has sometimes been supposed.     

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.     

Relative durations of conditioned stimulus and intertrial interval in conditioned suppression.

The effects of the relative durations of the conditional stimulus and the intertrial interval on bar pressing during a conditioned-suppression procedure were examined as a function of two additional variables--type of operant baseline schedule and rate of shock presentation. In Experiment 1, response suppression was compared across components of a multiple fixed-ratio, random-ratio, fixed-interval, random-interval schedule, at relative conditioned-stimulus/intertrial-interval durations of 1/1, 1/4, and 1/9. In Experiment 2, relative conditioned-stimulus/intertrial-interval duration (1/5, 3/3, or 5/1) was manipulated across groups, while shock frequency (2, 6, or 10 shocks/hr) was manipulated within groups. In both experiments, suppression during the signal was virtually complete at all relative durations. Responding was also suppressed during the intertrial interval, but that suppression varied as a function of experimental manipulations. In Experiment 1, intertrial-interval response rates were higher when relative signal duration was 1/9 than when it was 1/1, although both relative signal duration and shock frequency, which covaried, could have contributed to the difference. In Experiment 2, the patterning of response rates between successive shocks was affected by relative duration, absolute rates during the intertrial interval varied as a function of shock frequency, and differences between suppression during the signal and suppression during the intertrial interval were affected by both relative duration and shock frequency. The data support an analysis based upon relationships between shock-correlated and intertrial-interval stimuli and, as assessed by the relative-delay-to-reinforcement metric, are comparable to results that have been reported from experiments using similar manipulations under the autoshaping paradigm.     

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.     

Schedule-induced behavior in hyperactive children

A schedule-induced behavior paradigm was used to investigate the activity patterns of hyperactive children in a standardized situation. In Experiment I, 10 hyperactive and 10 normal control children matched for age, sex, and IQ were observed under conditions of baseline and schedule. Measures of a number of categorized activities were taken on a time-sampling basis. Hyperactive children were more active than controls in baseline and did not respond to the schedule, unlike the controls who became significantly more active in schedule conditions. In Experiment II, 12 hyperactive and 6 normal children were again subjected to the same experimental paradigm, but in two of the four experimental sessions the stimulant drug methylphenidate was administered in an attempt to reduce the amount of baseline activity. Results were substantially similar to those of Experiment I, with hyperactive children more active than controls in baseline and insensitive to the schedule. There was no overall effect of drug administration on the behavior of either group. There were some ratedependent effects of both drug and schedule conditions.     

SCHEDULE-INDUCED DRINKING: ELICITATION,ANTICIPATION, OR BEHAVIORAL INTERACTION?

We carried out five experiments with rats on fixed-time schedules in order to define the relation between drinking and individual food-pellet presentations. In Experiment 1, unsignaled extra food occurred at the end of occasional fixed intervals, and we compared subsequent drinking patterns with drinking before the extra food presentation. In Experiment 2 we presented signaled and unsignaled extra food and measured elicited and anticipatory drinking patterns. In Experiment 3, we observed the persistence of modified drinking patterns when several consecutive intervals ended with extra pellets. In Experiments 4 and 5, we varied the magnitude of food delivery across (rather than within) sessions to replicate published findings. Results show that schedule-induced drinking is neither elicited by food presentations nor induced by stimuli associated with a high food rate. All subjects seemed to follow a simple rule: during any stimulus signaling an increase in the local probability of food delivery within a session, engage in food-related behavior to the exclusion of drinking. Schedule-induced drinking appears to be the result of dynamic interactions among food-related behavior, drinking, and other motivated behavior, rather than a direct effect of the contingencies of food reinforcement.     

Drinking in a patchy environment: the effect of the price of water.

Rats in a laboratory foraging paradigm searched for sequential opportunities to drink in two water patches that differed in the bar-press price of each "sip" (20 licks) of water within a bout of drinking (Experiment 1) or the price and size (10, 20, or 40 licks) of each sip (Experiment 2). Total daily water intake was not affected by these variables. The rats responded faster at the patch where water was more costly. However, they accepted fewer opportunities to drink, and thus had fewer drinking bouts, and drinking bouts were smaller at the more costly patch than at the other patch. This resulted in the rats consuming a smaller proportion of their daily water from the more costly patch. The size of the differences in bout frequency and size between the patches appears to be based on the relative cost of water at the patches. The profitability of each patch was calculated in terms of the return (in milliliters) on either effort (bar presses) or time spent there. Although both measures were correlated with the relative total intake, bout size, and acceptance of opportunities at each patch, the time-based profitability was the better predictor of these intake measures. The rats did not minimize bar-press output; however, their choice between the patches and their bout sizes within patches varied in a way that reduced costs compared to what would have been expended drinking randomly. These data accord well with similar findings for choices among patches of food, suggesting that foraging for water and food occurs on the basis of comparable benefit-cost functions: In each case, the amount consumed is related to the time spent consuming.     

SOME RELATIONS BETWEEN CLASSICALLY CONDITIONED AGGRESSION AND CONDITIONED SUPPRESSION IN SQUIRREL MONKEYS

During three experiments with squirrel monkeys, stimulus and shock pairings were given in the presence of a bite tube. Experiments 1 and 2 used a conditioned-suppression procedure in which bar pressing was reinforced with food. A transparent shield prevented biting of the bar. When the stimulus was paired with shock, bar pressing decreased (conditioned suppression) and tube biting increased during the stimulus (classically conditioned aggression). When the bite tube was removed on alternate sessions in Experiment 2, there was more suppression when the tube was present, thus suggesting that biting competed with bar pressing. However, this simple competing-response interpretation was complicated by the findings of Experiment 3 where, with naive monkeys, bar pressing was never reinforced with food, yet bar pressing was induced during the stimulus and was highest when the bite tube was absent. The fact that stimulus-induced bar pressing developed indicates that bar pressing in conditioned-suppression procedures, suppressed or not, may be maintained by two types of control—the food reinforcer and induced CS control. The higher rate of induced bar pressing during the stimulus with the bite tube absent confounds a simple competing response interpretation of conditioned suppression. It suggests that shock-induced responses during conditioned suppression could be both contributing to and competing with responding maintained by food, with the net effect depending on specific but ill-defined features of the situation.     

Habituation in children within a behavior suppression paradigm

Using an instrumental response suppression paradigm, Experiment 1 demonstrated habituation and long-term retention of habituation to a tone stimulus in third-grade children, but specificity of habituation was not obtained when a new tone stimulus was presented. In Experiment 2 first graders showed specificity of habituation to auditory stimuli on the response suppression measure whereas fifth graders did not. However, both age groups evidenced specificity when a skin conductance measure was employed. In addition, specificity of habituation was not affected by amount of training at either age level. Experiment 3 demonstrated specificity of habituation in fifth graders on the response suppression measure when a cross-modality stimulus change was introduced. Implications of these data for analyses of habituation and traditional learning are discussed.