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.     

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.     

Separate neural sites for d-amphetamine suppression of mouse killing and feeding behavior in rats

The present study was conducted to investigate several possible neural sites for d-amphetamine's effect on mouse killing and feeding behaviors. d-Amphetamine (10, 20, and 30 μg) injected into each lateral ventricle, suppressed mouse kiling, food, and water intake in a dose-dependent manner. Bilateral adminstration of d-amphetamine (20 μg) into the central amygdaloid nucleus abolished mouse killing behavior but did not affect feeding and drinking. By contrast, bilateral amphetamine injections into the substantia nigra, or into the ventral region of the caudate nucleus, did not suppress mouse killing behavior, but significantly decreased food and water intake. The lateral hypothalamus was sensitive to d-amphetamine injections, which suppressed mouse killing and food intake as well as water intake. d-Amphetamine injections into the nucleus accumbens produced inconsistent effects on mouse killing and feeding. Our observations suggest a differentiation of the neural sites that mediate feeding from those underlying mouse killing behavior.     

Danger at the waterhole

The effects of declining accessibility of water and risk of electric shock on drinking patterns in rats were examined. Rats chose between two conditions to obtain their daily intake of water. In one condition, accessibility of water decreased systematically. In the other condition, water was readily accessible, but responses occasionally were followed by electric shock. Both variables affected drinking patterns similarly. As accessibility to water decreased, the number of drinking bouts initiated decreased but the quantity of water consumed increased. In response to risk of electric shock, the number of drinking bouts decreased but bout size increased. Results are considered in light of findings by Fanselow, Lester, and Helmstetter, (1988), Collier, Hirsch, and Hamlin (1972), and Marwine and Collier (1979).     

Interactive effects of deprivation in the albino rat

The interactive effects of feeding and drinking schedules were investigated in three experiments. Twenty-four hour water-deprived rats consumed their entire obligatory water intake prior to feeding, whereas 24-hr food-deprived rats consumed only small quantities of food prior to drinking. This drinking was apparently due to a shift of water stores rather than an actual negative water balance. Experiment 3 investigated the effects of 24, 48, or 72 hr of water, food, or total deprivation. Water-deprived rats did not adequately suppress food intake and became thirstier than totally deprived rats. The effects of total deprivation were essentially identical to those of food deprivation. These experiments indicate the degree to which deprivation schedules impose restrictions on the reinforcement parameters of behavioral experiments.     

DOES PACKAGE SIZE MATTER? A UNIT-PRICE ANALYSIS OF “DEMAND” FOR FOOD IN BABOONS

In a study examining “demand” for food, responding of 8 adult male baboons (Papio c. anubis) was maintained under a fixed-ratio schedule of food reinforcement during daily 23-hr experimental sessions. Completion of the ratio requirement resulted in the delivery of one, five, or 10 1-g food pellets. Supplemental feeding was limited to fruit and a dog biscuit daily. Responding increased as “cost” was increased across a wide range of fixed-ratio values before reaching a maximum and then decreasing. Increasing the number of food pellets per delivery decreased total responding and the number of reinforcements per day. A unit-price analysis, in which intake was converted to grams per day and fixed-ratio values were converted to responses per gram, yielded demand functions that overlapped at lower unit prices. Under one or more multiple-pellet conditions, however, intake decreased more quickly than under the one-pellet condition as the fixed-ratio value was increased in all but 1 baboon. This indicates that even when using unit-price conversions, there was variability in total intake. Although unit-price conversions yielded intake data that were more consistent across conditions, conditions differed in response topography even at the same unit prices: Under the multiple-pellet conditions there were longer pauses in responding, running response rate was slower, and the first eating bout (i.e., “meal”) of the session was smaller than under the one-pellet condition. These findings (a) support the heuristic value of a unit-price analysis for studying responding for and consumption of commodities that have similar attributes, and (b) indicate that different response topographies may result in similar intakes of a commodity.     

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.     

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.     

Within-session meal-size effects on induced drinking.

As a control for the effects of session duration and hunger on the relation between food magnitude and induced drinking, four food-deprived rats were exposed to a variable-time 50-s schedule of food delivery in which the size of each food delivery varied randomly within sessions. Food-related behavior and schedule-induced drinking per opportunity were examined as functions of meal size and postfood time. All rats showed an inverted-U-shaped relation between drinking per opportunity and meal size. This relation was caused by variation in the percentage of intervals that contained drinking and by variation in the number of drinking bouts per interval, rather than by bout duration or by the amount of drinking within those intervals that actually contained drinking. Head-in-feeder time increased linearly with meal size. Schedule-induced drinking was entrained by food delivery in 3 of 4 subjects; the entrainment was due to regulation of the starting time of each drinking bout rather than to regulation of bout duration.     

An economic analysis of "demand" for food in baboons.

Responding of 6 adult male baboons (Papio c. anubis) was maintained under a fixed-ratio schedule of food reinforcement during daily 22-hr experimental sessions. Completion of the ratio requirement resulted in the delivery of a single 1-g food pellet; supplemental feeding was limited to a daily fruit ration. Ratio values were increased on Mondays, Wednesdays, and Fridays according to the following schedule: 2, 4, 8, 16, 32, 64, 96, 128. Responding under each ratio value was examined four times. Under the Fixed-Ratio 2 conditions, food intake ranged between 300 and 600 g. Ratios were increased for each baboon until food intake decreased to about 100 g (20% to 30% of Fixed-Ratio 2 intake). Increasing the response cost increased total time responding and total daily responding in all baboons, but this increase in responding was not sufficient to maintain stable food intake. Baboons responded between 90 and 180 min per day. The highest running response rates were observed under the Fixed-Ratio 2 and Fixed-Ratio 4 schedules. Running rate was similar across the larger ratio values (greater than Fixed-Ratio 8) but was lower than that observed under the Fixed-Ratio 2 and Fixed-Ratio 4 schedules. Similar results were observed the four times that each fixed-ratio value was tested. Intake as a function of cost was analyzed by fitting data to the nonlinear equation proposed by Hursh, Raslear, Shurtleff, Bauman, and Simmons (1988) for "demand" functions. Demand for food was inelastic over most of the ratio values until food intake decreased to 15% to 55% of baseline. The results indicate that demand functions are appropriate for the study of food intake in baboons, but also caution that intake at the cost when demand shifts from inelastic to elastic and its relationship to maximal intake should also be included in analyses of demand for a commodity.     

Medial septal lesions: disruptions of microregulatory patterns and circadian rhythmicity in rats

The microregulatory patterns of food and water intake were examined in male and female rats bearing medical septal lesions and in sham-operated controls. Medial septal ablation, although not affecting the total amount of food or water ingested, resulted in a profound disruption of the pattern of intake. Circadian rhythmicity was disrupted for a period, returning to normal by 25 days postlesion. Permanent disruptions occurred in feeding patterns in the rats with septal lesions ingested more frequent but smaller meals. There was also a marked increase in food-intake-associated drinking and a decrease in non-food-intake-associated drinking. The results are interpreted to reflect two separate independent effects, a general circadian disruption and an alteration in requlatory behavior produced by a chronic depletion of body fluid.     

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.     

Schedule-induced polydipsia in rats living in an operant environment

The effect of variations in interreinforcement interval on the temporal and distributional relation between feeding and drinking was continuously monitored. Rats were housed continuously in an operant chamber in which water was freely available, but lever pressing was required to obtain food (45-mg pellets). Initially, pellets were delivered on a fixed-ratio 1 schedule of reinforcement, which was followed by testing on response-initiated fixed-interval 15-, 30-, and 60-second schedules. The total number of discrete, daily meals (a period in which several pellets were earned in succession) was slightly higher during the fixed-interval schedules than during the fixed-ratio 1, but there was no systematic effect of fixed-interval length on meal frequency. Total water consumption, in contrast, increased dramatically as the interval was lengthened: both subjects consumed two to three times as much water on the fixed-interval 60-second schedule as on the fixed-ratio 1. The increased water consumption was the result of an alteration in the distribution of drinking relative to eating. During the fixed-ratio 1 condition, drinking occurred infrequently following individual food pellets and represented the smallest percentage of total drinking; drinking occurred predominantly just before or after a meal. As the fixed interval was lengthened, however, the frequency of postpellet drinking gradually increased and eventually comprised the largest proportion of daily drinking.     

Effects of lick-contingent timeout on schedule-induced polydipsia

Rats bar pressing on a 1-min fixed-interval schedule for 45-mg food pellets became polydipsic when water was concurrently available. They were then exposed to conditions in which each lick on the drinking tube produced a timeout period during which the food-schedule lever was retracted and the fixed-interval timer either did or did not continue to operate. Licks occurring within a timeout period extended its duration. As the duration of the lick-initiated timeout period was increased logarithmically through four values from 10 sec to 80 sec, lick rates as well as water intake rates generally decreased for all three subjects. As timeout duration was progressively increased, the rate of licks occurring in the absense of, but producing, timeouts decreased for all three rats, whereas the rate of licks occurring in the presence of timeout periods remained essentially constant. Water-intake rates and, with one exception, lick rates were suppressed more by timeout periods during which the fixed-interval timer did not continue to operate. These results indicate that lick-contingent timeout from positive reinforcement reduces schedule-induced drinking, and this suppressive effect is greater when the timeout period necessarily increases the interreinforcement interval beyond its minimum duration than when it does not.     

Inhibition of ad Libitum Feeding in Rats by Salt Injections and Water Deprivation

Inhibition of ad libitum feeding in rats was induced by hypertonic NaCl injections. Though osmotic loads of sufficient size were capable of abolishing feeding completely for a time, the effect was not as large as had been predicted from a hypothesis of strictly linear subtractive inhibition. Feeding at a low level of hunger seems to be somewhat less affected by osmotic inhibition than feeding on a deprivation schedule. Inhibition of feeding was also produced by deprivation of water, and both the inhibition of food intake during deprivation, and the disinhibition by subsequent drinking indicated that the amount of inhibition of food intake is a non-linear (accelerating) function of water deficit. A model of the process indicating that the thirst signal undergoes a non-linear transformation before being subtracted from the signal corresponding to food demand is proposed.     

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.     

Four effects of exogenous insulin on food intake

After a subcutaneous injection of bovine insulin into the rat, at first there is an augmentation of the satiety produced by nutrient eaten immediately before injection. Later, with large enough doses, as has been commonly observed, feeding is elicited—perhaps simply by hastening the passage of satiety. A third type of effect is behavioural disruption, reducing food and water intake when food is withheld for an hour after injection and producing postural changes even when food is present. Fourth, repeated pairing of insulin injection with intake of water of a particular flavour (even when drunk over half an hour beforehand) depresses subsequent intake of water having that flavour, whether presented alone or together with water of another flavour which has been paired with control injections. The acquired discriminated intake change involves the initial acceptability of the flavour but changes in the inhibition of acceptability during an intake bout have not been excluded.     

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.     

Motivation in concurrent variable-interval schedules with food and water reinforcers

The lever pressing of four food- and water-deprived rats was reinforced on concurrent variable-interval schedules. Food reinforced one response, and water reinforced the other. Response rates in baseline were higher in the food component than in the water component. After response patterns and body weights had stabilized, the animals were given access to either food only, water only, both food and water, or neither food nor water (baseline) before daily sessions. Giving food before a session decreased per cent time in the food component, decreased overall response rates for food, and increased overall response rates for water. Giving water before a session increased per cent time in the food component, increased overall response rates for food, and decreased overall response rates for water. Giving both food and water before a session resulted in a combination of prefeeding and prewatering effects. More food and more water were consumed when both were available than when only one was available before a session.