Dynamic effects of food magnitude on interim-terminal interaction

We tested the assumption of a facilitatory relation between periodic food presentation and schedule-induced drinking by examination of (a) elicited drinking, (b) drinking in anticipation of food delivery, and (c) possible indirect effects of food delivery on drinking. We exposed rats to a fixed-time 60-second schedule in which interfood intervals ended in either one or four food pellets with equal probability. In Phases 1 and 3, a stimulus signaled the magnitude of upcoming food presentation. In Phase 2, the stimulus was eliminated. Changes in drinking and “head-in-feeder” distributions within interfood intervals demonstrated that head-in-feeder was controlled directly by food presentation, but drinking was not. Head-in-feeder increased and drinking was reduced when large meals began or ended an interval. In Phases 4 to 6, meal size was manipulated across sessions yielding a positive relation between meal size and schedule-induced drinking. We conclude: (1) Schedule-induced drinking is determined by distributions of food-related behavior and results from indirect effects of food delivery; and (2) the amount of schedule-induced drinking and the form of the drinking distributions in this experiment can be accurately explained by two assumptions: (a) Food presentation facilitates food-related behavior through elicitation and anticipation; and (b) food-related behavior and drinking are reciprocally, linearly related.     

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.     

Schedule-induced drinking as a function of percentage reinforcement

Drinking was recorded in rats while lever pressing was maintained on a series of percentage reinforcement schedules in which the per cent of 1-min fixed intervals terminating with food was 100, 90, 30, 70, 10, 50, and 100%. Intervals in which a pellet was omitted were terminated by brief light flash and click stimuli that were also correlated with food presentations. Drinking failed to develop in five of six subjects following intervals in which the brief stimuli were presented regardless of percentage reinforcement. Postpellet drinking, which followed intervals terminated with pellet delivery, however, increased in both duration and amount ingested per interval as percentage reinforcement was systematically decreased. The increase in postpellet drinking above that produced by 100% reinforcement was interpreted as an analogue of the positive-contrast effect observed with food-reinforced operants.     

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.     

Abdominal vagotomy disrupts food-related drinking in the rat

Rats with complete subdiaphragmatic bilateral transection of the abdominal vagus (Vgx-C) showed disordered food-related drinking when drinking water in temporal association with a meal of dry food after 5-hr food deprivation and when drinking water in association with a liquid meal after 24-hr food deprivation. The Vgx-C rats drank after significantly longer latencies and drank significantly less water in 1 hr than did sham-vagotomized (Sham) rats after eating the same size meal (solid or liquid) as Shams. Rats with incomplete vagal transection (Vgx-I) ate and drank like Shams. Water intake of Sham and Vgx-I rats correlated positively with the meal size of solid food, but the water intake of Vgx-C rats did not. The failure of Vgx-C rats to drink water normally when food was ingested was not due to failure of a food stimulus to reach the intestine, because Vgx-C and Sham rats emptied equivalent volumes of liquid food from the stomach into the intestine within 10 min of food entering the stomach. These results indicate that the abdominal vagus is an important neurological substrate for food-related drinking in the rat.     

Schedule-induced drinking as functions of interpellet interval and draught size in the Java macaque

Three Java monkeys received food pellets that were assigned by both ascending and descending series of fixed-time schedules whose values varied between 8 and 256 seconds. The draught size dispensed by a concurrently available water-delivery tube was systematically varied between 1.0 and 0.3 milliliter per lick at various fixed-time values during the second and third series determinations. Session water intake was bitonically related to the interpellet interval and was determined by the interaction of (1) the probability of initiating a drinking bout, which fell off at the highest interpellet intervals and, (2) the size of the bout, which increased directly with increases in interpellet interval. Variations in draught size had little effect on total session intakes, but reduced bout size at draught sizes of 0.5 milliliter and below. Thus, a volume-regulation process of schedule-induced drinking operated generally at the session-intake level, but was limited to higher draught sizes at the bout level.     

Food and water intake as functions of resource consumption costs in a closed economy.

In two experiments, rats living in a closed economy were offered continuous, concurrent access to four resources: food, water, a nest, and a running wheel. Costs of consuming food and water were imposed with bar-press requirements, and the price of either one or both resources was raised. As the consumption cost increased, less was consumed in each bout of resource use. Bout frequency increased, but not sufficiently to compensate for the fall in bout size, and total intake fell. Food and water tended to be complementary resources, in that as intake of one fell with its price, intake of the other also decreased. This interaction was accounted for by the defense of the ratio of body water to lean body mass. As amount consumed decreased, increases in feed efficiency (weight gain per unit of food ingested) and the use of stored calories compensated for the reduced energy intake. There was evidence of competition between feeding and drinking at the higher costs: When both commodities were expensive, the decline in the intake of each one was greater than when only one commodity was expensive. Although the time spent nesting, running, and in unmonitored activity was adjusted when feeding or drinking took more of the rat's day, there was no particular activity that was sacrificed.     

Schedule-induced drinking as a function of interreinforcement interval in the rhesus monkey

Lever presses by two rhesus monkeys produced food pellets that were assigned by both an ascending and descending series of fixed-interval schedules whose values varied between 1 and 512 sec. The amount of schedule-induced drinking was bitonically related to interreinforcement interval, reaching a maximum at approximately 120 sec and declining at longer fixed intervals. The relation between water intake and interreinforcement interval was complexly related to two drinking measures: (1) the probability of drinking following a pellet and (2) the amount drunk per bout. Drinking rate was also bitonically related to interreinforcement interval.     

SCHEDULE-INDUCED POLYDIPSIA: ARE RESPONSE-DEPENDENT SCHEDULES A LIMITING CONDITION?1

The collateral water intake of albino rats was measured when the inter-pellet intervals in fixed-ratio and fixed-time schedules were equated. Fixed-ratio and fixed-time inter-pellet intervals were equated by dividing the average fixed-ratio session time of each subject by 150 (food pellets). The average inter-pellet interval obtained then defined the subsequent fixed-time schedule for each individual subject. Shifts to fixed-time schedules followed the completion of each fixed-ratio 20, 40, and 80 schedule. This procedure permitted an assessment of the extent to which excessive collateral drinking was associated with inter-pellet interval length or adventitious food reinforcement. For both the fixed-ratio and fixed-time schedules, drinking progressively increased as a function of increasing the duration of the inter-pellet interval and was a post-pellet event under the control of variables other than adventitious food reinforcement.     

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.     

An analysis of rats' drinking-tube contacts under tandem and fixed-interval schedules of food presentation

Rats' lever presses and drinking-tube contacts were studied under fixed-interval schedules of food presentation and under a tandem schedule composed of three fixed intervals. One group of rats was exposed first to the tandem schedule, next to fixed-interval schedules of comparable interpellet intervals, and once again to the tandem schedule; a second group of rats was exposed first to a fixed-interval and then to the tandem schedule. Under the tandem schedule, lever presses occurred at a higher rate and were more uniformly distributed in time than under the fixed-interval schedule. Tube contacts emitted by rats exposed first to a fixed-interval schedule consisted mostly of tongue contacts, which occurred at a high rate shortly after food; tube contacts emitted by rats exposed first to the tandem schedule consisted mostly of paw contacts, which occurred at a lower rate at times other than shortly after food. Changing the schedule from fixed interval to tandem decreased the frequency of tongue contacts for all rats. Under schedules of food presentation with comparable interpellet intervals, the schedule of food presentation, rather than the rate of food delivery per se, determined the topography and temporal locus of drinking-tube contacts.     

Successive interresponse times in fixed-ratio and second-order fixed-ratio performance

Three rats were trained on a schedule in which every sixth response produced a timeout of 5 sec minimum duration, and food was delivered at the onset of timeout. Successive interresponse times were measured under these conditions, and also when behavior was maintained by second-order fixed-ratio and fixed-interval schedules. Under the second-order schedules, each six-response fixed-ratio component was followed by a timeout, and occasionally food was delivered at the onset of a timeout. In the fixed-ratio schedule, the successive interresponse times showed a decrease followed by an increase before food delivery, but this systematic variation in interresponse times was not found when the performance was under second-order reinforcement. Under both second-order schedules the latencies of successive components, and the successive interresponse times within each component, showed a decrease as food delivery was approached.     

Brief successive temporal observational sampling as a possible indicator of daily overt seizure activity in epileptic rats

Chronic, limbic epilepsy was induced in male rats by a single systemic injection of lithium and pilocarpine. In two separate experiments, each 6 mo. in duration, the proportion of the population that displayed an overt motor seizure (head nodding, rearing, and forelimb clonus) within 10 min. each day after delivery of food pellets (despite food provided ad libitum) was recorded. Cumulative records for the numbers of rats in each population that displayed overt seizure for the first time during the observation period were plotted as a function of time. The occurrence of the saturated (approaching 100%) asymptote at about five months was interpreted to indicate that brief (10 min.) sampling over the 144 successive days that would compose a single 24-hr. period could serve as an estimate of the average probability of an overt seizure during an average 24-hr. period. If this assumption is correct, then these rats would have displayed about one overt seizure per day.     

The reinforcement value of schedule-induced drinking

The effect of food reinforcement schedules on the reinforcement value of drinking water was evaluated. Food-deprived rats were exposed to concurrent, identical variable-time schedules of food presentation, the food thus being delivered independently of the rats' behavior. When the relative amount of time spent in a schedule component stabilized, an opportunity to drink water was introduced into one schedule component. The value of the variable-time schedules was varied from 60 to 90 to 270 sec. The relative amount of time spent in the schedule component associated with drinking water was a decreasing function of food frequency for two animals and remained constant for the third. Drinking rates were direct functions of food frequency, and the amount of water drunk per pellet was an inverse function of food frequency. The reinforcement value of drinking water, according to the Matching Law, was a direct function of the frequency of food presentation. It was concluded that food reinforcement schedules indirectly influence rates of drinking by altering the reinforcement value of drinking water and that certain properties of schedule-induced drinking can be accounted for in terms of the reinforcement value of drinking water, the rate of drinking, and the frequency of food presentation.     

The economics of the law of effect.

A corollary of the law of effect predicts that the larger the reinforcement, the greater the rate of responding. However, an animal must eat more small portions than large portions to obtain the same daily intake, and one would predict, therefore, that when eating smaller portions an efficient animal would eat less (conserving time and energy) and/or respond faster (conserving time). The latter of these predictions was supported by the present experiments with free-feeding rats for which portion size (pellet size or duration of feeder presentation) and portion price within meals were varied. Response rate was a function of the unit price (responses/g) of food: Rats responded faster when portions were smaller or when prices were higher. Meal size and frequency were relatively unaffected by unit price, but were influenced by the price of meal initiation. The results are discussed in relation to the economic differences between traditional operant and free-feeding paradigms and to both traditional and more recent formulations of the law of effect.     

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.     

Polydipsia induced in the rat by a second-order schedule

Drinking was studied in rats pressing a bar on a second-order schedule in which every third completion of a 1-min fixed interval was followed by food presentation. A brief flash of light signaled the completion of each fixed-interval component. The rats drank not only after the food presentations but also after presentations of the light flash alone. A high rate of steady drinking followed intervals terminated by a food presentation. Drinking that followed intervals terminated by a light flash alone was of comparable rate, but characteristically interrupted by bar pressing. When 250-mg food pellets were used instead of 45-mg pellets, both drinking and bar-pressing rates increased substantially.     

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.     

Eating and drinking: An economic analysis

Food-deprived rats were exposed to various schedules of food delivery; water-deprived rats were exposed to various schedules of water delivery. Eating and drinking were measured over sessions and at points throughout sessions. The symmetries and asymmetries of food and water consumption were explored in terms of: (1) substitutability of food versus water, and of food and water on the one hand versus leisure on the other, (2) constraints imposed by various schedules of food and water, and (3) the tendency of rats to maximize utility within the imposed constraints.