Meal patterns of cats encountering variable food procurement cost.

The meal patterns of 2 cats in a laboratory habitat with variable foraging costs were examined in a foraging paradigm in which subjects could initiate meals at any time by completing a predetermined number of bar presses (the procurement price) and then could eat any amount. From meal to meal, the procurement price either was fixed or varied among a geometric series of five prices. As the fixed price or the mean of the variable prices increased, meal frequency decreased and meal size increased; daily intake was unaffected. Within variable-price schedules, meal size was not related to the just-paid procurement price. These results suggest that cats respond to the global rather than to the local cost structure of their habitat. They appear to respond to an average of the prices encountered, initiating meals of a frequency and size appropriate to that average. This was true even when the average price was high, meals were infrequent, and thus price encounters were widely separated in time. Therefore, the time window over which the consequences of behavior can affect behavior is longer than often conceived, at least in economies in which the animal controls its intake and the frequency, size, and distribution of its meals.     

Meal patterns following changes in procurement cost for rats with fornix transection

The purpose of the present study was to explore the effects of a varied procurement cost on the foraging behavior of rats with fornix transection. An operant analog of foraging requirements was used to examine the feeding patterns of the animals under free feeding, low procurement cost (FR5), and high procurement cost (FR80) situations, in an environment with minimal sensory distraction. It was found that animals with fornix transection did not differ from control rats in general consumption. Both groups were also able to adapt their feeding behavior to the varied procurement cost. As the procurement cost increased, the number of meals consumed decreased while the meal duration increased. The meal patterns themselves were different for the fornix transected animals and the control group. Animals with fornix transections ate more meals over the course of a day than did control animals; their meals were of a longer duration, and their intermeal intervals were shorter than those of control animals. During the course of a meal, the rats with fornix transections took a larger number of breaks, during which they drank, explored, or engaged in activities other than eating. These differences in the feeding patterns were seen across all procurement cost levels. The data support the possibility of hippocampal involvement in behavioral organization or sequencing.     

Reinforcement of inhibition

A differential-reinforcement-of-other-behavior (DRO) schedule with trials and delayed reinforcement was investigated. Periodically a wheel was briefly available to rats, followed six seconds later by brief availability of a bar. Variable-ratio food reinforcement of wheel turns was adjusted to give 95% turns. After variable-ratio-five reinforcement of bar presses produced 100% pressing, then separate ratio schedules were used for presses following turns (turn presses) and presses following nonturns (nonturn presses). Increasing nonturn-press reinforcements decreased turns, even though total reinforcements increased. Reversal by decreasing nonturn-press reinforcements raised turns, though with hysteresis. Thus food reinforcement increased nonturns even though delayed six to ten seconds after nonturns, a delay that greatly reduces response reinforcement. Those and other results indicate that the turn decrease was not due to reinforcement of competing responses. Evidence against other alternatives, and the reduction of responding by increased reinforcement, indicate that the term inhibition is appropriate for the phenomenon reinforced. Response-specific inhibition appears appropriate for this particular kind, since its effects are more specific to particular responses than Pavlovian conditioned-inhibition. Response-specific inhibition seems best considered a behavioral output comparable to responses (e.g., both reinforcible) but with important properties different from responses (e.g., different reinforcement-delay gradients).     

Procurement time as a determinant of meal frequency and meal duration.

Foraging involves the expenditure of both time and effort in the acquisition of food; animals typically modify their meal patterns so as to reduce these expenditures or costs. The contribution of time, as compared with effort, to the overall cost perceived by an animal is not known. We investigated the effect of foraging time as a cost independent of effort by measuring the meal patterns of rats living in a laboratory foraging simulation in which they earned all their daily intake. They pressed a bar once to initiate an interval (procurement interval) leading to the presentation of a large cup of food from which they could eat a meal of any size. As the length of the interval increased from 1 s to 46 hr, meal frequency decreased regularly. Meal size increased in a compensatory fashion, and total daily intake was conserved through an interval of 23 hr. The changes in meal frequency occurred because of changes in the rat's latency to bar press after each meal. The functions relating meal frequency and size to the procurement interval were of the same shape as those seen when cost is the completion of a bar-press requirement, which entails the expenditure of both effort and time. When the bar-press requirement was increased to 10, meal frequency was reduced, but time and effort did not appear to simply add together in the rat's perception of cost. These data reveal that time is preceived to be a cost by rats foraging in this laboratory environment. These results suggest that the time parameters of foraging are different from those of consumption.     

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.     

Preadaptation to the feeding schedule does not eliminate activity-based anorexia in rats

To test whether activity-based anorexia (ABA) still occurs after preadaptation to the feeding schedule, 20 rats were first exposed to a feeding schedule of one 90-min meal per day until adaptation occurred (measured by maintenance of stable body weight). Then, during ABA training, half the rats (wheel group) were confined in running wheels except during the daily meal, and half (cage group) were not. Wheel running suppressed feeding—that is, food intake in the wheel group was less than that in the cage group. Also, the rats in the wheel group lost weight, whereas those in the cage group did not. Wheel running increased over days. Thus, the defining characteristics of ABA were evident in rats that were not subjected to ABA training until after they had become well adapted to the feeding schedule. These findings support the view that the suppression of feeding produced by wheel running triggers the vicious circle of ABA. They also cast doubt on the hypothesis that activity-induced interference with adaptation to the feeding schedule plays a key role in causing ABA.     

Deprivation and satiation: The interrelations between food and wheel running

Two experiments were designed to assess whether depriving rats of food would increase the reinforcement effectiveness of wheel running (Experiment 1) and whether satiation for wheel running would decrease the reinforcement effectiveness of food (Experiment 2). In Experiment 1, a progressive-ratio schedule was used to measure the reinforcement effectiveness of wheel running when rats were deprived or not deprived of food. Completion of a fixed number of lever presses released a brake on a running wheel for 60 s, and the response requirement was systematically increased until the rat stopped pressing or until 8 hr had elapsed. The ratio value reached (and the total number of lever presses) was an inverted-U function of food deprivation (percentage body weight). In Experiment 2, when wheel running preceded test sessions, fewer food-reinforced lever presses were maintained by the progressive-ratio schedule, and responding occurred at a lower rate on a variable-interval schedule. An interpretation of these results is that deprivation or satiation with respect to one event (such as food) alters the reinforcement effectiveness of a different event (such as access to wheel running).     

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.     

Transfer of effort across behaviors

A series of experiments investigated the effects of required force and number of responses per reinforcer upon the subsequent performance of a second behavior. In the first experiment, a group of rats required to complete five round trips in an alley per food pellet subsequently bar-pressed for food at a greater rate than a group rewarded for each round trip or a control group that did not receive the alley experience. In the second experiment, a group required to apply a 70-g bar-press force subsequently shuttled for food at a greater rate than a group required merely to touch the lever or a control group that did not undergo the lever-press manipulation. The third experiment found that the force effect persisted across all five test sessions and was attributable to differences both in response speed and interresponse time. The fourth experiment found that both the necessary bar-press force and number of bar presses per reward affected subsequent shuttling in extinction. Two alternative interpretations of these results were compared: (a) the degree of accustomed effort per reinforcer becomes a generalized component of instrumental behavior or (b) high effort increases the habituation frustration-produced disruptive responses.     

Changes in feeding and foraging patterns as an antipredator defensive strategy: a laboratory simulation using aversive stimulation in a closed economy.

The effects of the risk of electric shock on the meal patterns of rats living in an operant chamber were investigated. Rats could obtain food by working on a response lever that provided reinforcement according to chained fixed-ratio continuous reinforcement schedules that allowed the animals control over meal size. Using a two-compartment operant chamber with a safe nesting area and manipulanda area with a grid floor, shock could be correlated with responding on the schedule. Shocks (less than or equal to 1.25 per hour) were scheduled to occur randomly throughout the day, independent of the rat's behavior. Shock caused a reorganization of meal patterns such that the animals took less frequent but larger meals. This pattern reduced the time the animals spent at risk without compromising caloric balance. Similar changes in feeding pattern were obtained in both hooded and albino rats. Exposure to shock in a separate chamber did not produce these behavioral modifications. The magnitude of shock-induced alterations of meal patterns was greater with chained fixed-ratio 90 continuous reinforcement than with chained fixed-ratio 10 continuous reinforcement. Additionally, the rats seemed to be able to reduce food intake but increase caloric efficiency, such that the reduced food intake did not have deleterious effects on maintenance of body weight. These behavioral modifications reduced the number of shocks received from that which would have been expected if meal pattern changes had not occurred. We suggest that this technique may provide a useful laboratory simulation of the impact that the risk of predation has on foraging behavior.     

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.     

Generation of different numbers of button presses under supplementary feedback conditions by educable retardates.

Educable retardates (10 boys and 10 girls) generated varying numbers of presses on a response key under 5 different feedback conditions. Neither time per response nor accuracy was affected by feedback condition, but time per response and magnitude of errors increased when larger numbers of responses were required.     

Economic substitutability of electrical brain stimulation, food, and water.

Concurrent variable-ratio schedules of electrical brain stimulation, food, and water were paired in various combinations as reinforcement of rats' lever presses. Relative prices of the concurrent reinforcers were varied by changing the ratio of the response requirements on the two levers. Economic substitutability, measured by the sensitivity of response ratio to changes in relative price, was highest with brain stimulation reinforcement of presses on both levers and lowest with food reinforcement of presses on one lever and water reinforcement of presses on the other. Substitutability with brain stimulation reinforcement of presses on one lever and either food or water reinforcement for presses on the other was about as high as with brain stimulation for presses on both levers. Electrical brain stimulation for rats may thus serve as an economic substitute for two reinforcers, neither of which is substitutable for the other.     

Ontogeny of meal patterning in rats and its recapitulation during recovery from lateral hypothalamic lesions.

The development of feeding patterns was investigated in weanling rats and in rats with lateral hypothalamic lesions. From 16 to 25 days of age, the weanlings demonstrated a preprandial intake pattern, i.e., a positive correlation between meal size and time since the preceding meal. This subsequently declined while the postprandial relationship (correlation with time until subsequent meal) began to emerge such that by 30-35 days of age a full adult pattern was observed. Rats recovering from lateral hypothalamic lesions, for a brief period, also demonstrated a preprandial intake pattern. The postprandial relationship was abolished by the lesions. These results suggest that the development of adult meal patterning results from maturation of lateral hypothalamic mechanisms governing meal initiation.     

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

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

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.     

Substitution and caloric regulation in a closed economy.

Three experiments were conducted to study the effect of an imperfect substitute for food on demand for food in a closed economy. In Experiments 1 and 2, rats pressed a lever for their entire daily food ration, and a fixed ratio of presses was required for each food pellet. In both experiments, the fixed ratio was held constant during a daily session but was increased between sessions. The fixed ratio was increased over a series of daily sessions once in the absence of concurrently available sucrose and again when sucrose pellets were freely available. For both series, increases in the fixed ratio reduced food intake, but body weight was reduced only in the no-sucrose condition. In the sucrose condition, body weight and total caloric intake (sucrose plus food) were relatively unaffected by increases in the fixed ratio. At all fixed ratios, food intake was proportionally reduced by the intake of sucrose. In Experiment 3, monkeys obtained food or saccharin by pressing keys; the fixed ratio of presses per food pellet was increased once when tap water was each monkey's only source of fluid, again when each monkey's water was sweetened with saccharin, and a third time when each monkey had concurrent access to the saccharin solution and plain water. Increases in the fixed ratio, but not the intake of the saccharin solution, reduced each monkey's food intake. Because neither rats' sucrose nor monkeys' saccharin intakes affected the slope of the respective demand curves for food, monkeys and rats increased their daily output of presses and thereby defended their daily intake of those complementary elements of food. However, sucrose reduced rats' food intake. The relative constancy of body weight and total caloric intake in the sucrose condition is consistent with the possibility that rats tended to regulate caloric intake.     

Low-response-rate conditioning history and fixed-interval responding in rats.

Bar presses by one group of rats were conditioned under a differential-reinforcement-of-low-rate reinforcement schedule immediately prior to conditioning under a fixed-interval schedule. In a second group of rats, bar presses were conditioned first under a differential-reinforcement-of-low-rate schedule and then under a fixed-ratio schedule prior to conditioning under a fixed-interval schedule. Low response rates occurred under the fixed-interval schedule only when it was immediately preceded by low-rate conditioning. Otherwise, fixed-interval responding was similar to responding under the fixed-ratio schedule. This finding suggests that responses of laboratory animals are sensitive to immediate history, and, unlike human responses, are relatively insensitive to a history of low-rate conditioning when it is followed by high-rate conditioning.     

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.     

Rats' lever-press durations as psychophysical judgments of time

Hungry rats received food following lever-press durations exceeding a minimum value, which ranged from 0 to 6.4 sec. When no intertrial intervals separated successive presses, modal press durations remained at very short values as the minimum value required for food was increased. This was particularly true immediately after a food presentation. When an 8-sec intertrial interval followed each lever release, modal press durations were always at or beyond the minimum value required for food, and outcome of the preceding press had no effect on press duration. Possible reasons for the effects of intertrial intervals included punishment of short presses, increased delay of reinforcement of short presses, and reduced density of reinforcement. In addition, functions relating discrete-trials lever-press duration to minimum duration required for food were found to be qualitatively and quantitatively similar to the power functions recently proposed by Catania (1970) for interresponse time and response latency. This similarity was taken as support for a general psychophysical law of temporal judgments.