Time horizons in rats: the effect of operant control of access to future food.

The primary goal of this experiment was to determine whether the addition of an operant requirement for access to a less costly (continuous reinforcement) patch of future food increased the time horizon over which that future patch decreased intake in a currently available depleting (progressive-ratio) patch. Three groups of 4 rats were tested. Each member of the earned-time group was required to cumulate a fixed-time outside the progressive-ratio patch to obtain access to food in the less costly patch; the fixed-time requirement ranged from 2 to 64 min. Rats in the matched-time group received response-independent access to less costly food at the average delay shown by the earned-time group. Rats in the matched-time no-food group were removed from the chamber at the same average delay without receiving access to less costly food. Two of the earned-time rats showed an increased time horizon relative to that shown by the matched-time rats (approaching 40 min for 1 rat). The other 2 earned-time rats markedly increased instrumental responding but showed suppression of intake only when food was less than 20 min away. The matched-time group showed less suppression of intake over a similar range of delay intervals. Surprisingly, the matched-time no-food animals also showed suppression of intake concentrated at the end of the session, possibly reflecting the receipt of their entire daily ration 30 min after the session. The potential importance of time horizons to the foraging process is clear, but experimenters are still working out paradigms for investigation of these horizons.     

Effects of anorectic drugs on food intake under progressive-ratio and free-access conditions in rats

The effects of two anorectic drugs, dexfenfluramine and phentermine, on food intake under different food-access conditions were examined. Experiment 1 compared the effects of these drugs on food intake under a progressive-ratio (PR) schedule and free-access conditions. Dexfenfluramine decreased food intake under both conditions, but the doses required to decrease intake under free-access conditions were higher than those required to reduce intake under the PR condition. Intermediate doses of phentermine sometimes increased breaking points, and higher doses decreased them. Phentermine decreased food intake at the same doses under both access conditions. Thus the potency of dexfenfluramine, but not phentermine, to decrease food-maintained behavior depended upon the food-access condition. Experiment 2 used a novel mixed progressive-ratio schedule of food delivery to study the duration of drug effects. Sessions consisted of five components separated by 3-hr timeouts. The ratio requirement reset at the beginning of each component and a new breaking point was obtained. Both dexfenfluramine and phentermine dose-dependently decreased breaking points early in the session. In some rats, compensatory increases in breaking point were observed. That is, breaking points later in the session increased over control levels, resulting in no change in the total number of food pellets earned for the session compared to control. The present findings suggest that the effects of some anorectic drugs depend upon the access conditions for food; increasing the effort to obtain food may enhance their ability to decrease food-maintained behavior.     

Techniques for establishing schedules with wheel running as reinforcement in rats.

In three experiments, access to wheel running was contingent on lever pressing. In each experiment, the duration of access to running was reduced gradually to 4, 5, or 6 s, and the schedule parameters were expanded gradually. The sessions lasted 2 hr. In Experiment 1, a fixed-ratio 20 schedule controlled a typical break-and-run pattern of lever pressing that was maintained throughout the session for 3 rats. In Experiment 2, a fixed-interval schedule of 6 min maintained lever pressing throughout the session for 3 rats, and for 1 rat, the rate of lever pressing was positively accelerated between reinforcements. In Experiment 3, a variable-ratio schedule of 20 or 35 was in effect and maintained lever pressing at a very stable pace throughout the session for 2 of 3 rats; for 1 rat, lever pressing was maintained at an irregular rate. When the session duration was extended to successive 24-hr periods, with food and water accessible in Experiment 3, lever pressing settled into a periodic pattern occurring at a high rate at approximately the same time each day. In each experiment, the rats that developed the highest local rates of running during wheel access also maintained the most stable and highest rates of lever pressing.     

Assessment of attack and drinking in White King pigeons on response-independent food schedules.

Four White King pigeons in Experiment I were exposed to a fixed-time 90-second food schedule with successive access to water and a conspecific target. Drinking per session was sporadic and minimal, while attack per session occurred during most interfood intervals for all animals. Analysis of the temporal distribution of attack showed that the typical postreinforcement pattern of attack developed over the course of the experiment. In Experiment II, the same animals were exposed to a series of fixed-time schedules ranging from 30 to 360 seconds with successive access to water and target. Time engaged in drinking and the number of interfood intervals with drinking were less than that of attack. Food and no-food baselines, which have been typically used to assess schedules-induced drinking and attack, respectively, were used to evaluate the effect of the schedule on attack and water ingestion. Relative to the no-food baseline, both attack and drinking were enhanced by the schedule in all birds. Relative to the food baseline, drinking was slightly suppressed in three birds and attack was enhanced in all. For all animals, the food baseline resulted in more attack and drinking than the no-food baseline.     

Time horizons in rats foraging for food in temporally separated patches

An important tenet of optimal foraging theory is that foragers compare prey densities in alternative patches to determine an optimal distribution of foraging behavior over time. A critical question is over what time period (time horizon) this integration of information and behavior occurs. Recent research has indicated that rats do not compare food density in a depleting patch with that in a rich patch delayed by an hour or more (Timberlake, 1984). In the present research we attempted to specify over what time period a future rich patch would affect current foraging. The effect of future food was measured by early entry into the rich patch (anticipation) and by a decrease in food obtained in the depleting patch (suppression). The rats showed anticipation of a rich patch up to an hour distant, but suppressed current feeding only if the rich patch was 16 min distant or less. The suppression effect appeared mediated by competition for expression between anticipatory entries into the rich patch and continued foraging in the depleting patch. These results suggest that optimal foraging is based on a variety of specific mechanisms rather than a general optimizing algorithm with a single time horizon.     

Food and amphetamine self-administration by baboons: effects of alternatives.

The effects of the availability of an alternative reinforcer on responding maintained by food pellets or fluid solutions were examined in 6 adult male baboons (Papio cynocephalus anubis). During daily 23-hr experimental sessions, baboons had concurrent access to both food pellets and fluid, with responding maintained under fixed-ratio schedules of reinforcement that varied between the two commodities. The fixed-ratio requirement, or cost, for pellets was increased when (a) no fluid, (b) a dilute dextrose vehicle, (c) 0.002 mg/kg d-amphetamine, or (d) 0.004 mg/kg d-amphetamine was available. When given nonrestricted concurrent access to food pellets and amphetamine at minimal cost (FR 2), baboons self-administered sufficient amphetamine to decrease pellet intake. Increasing the response requirement for pellets decreased pellet intake at a similar rate regardless of the available fluid and increased fluid intake in a variable manner among baboons such that there were no statistically significant increases in fluid intake. In contrast, when access to pellets was restricted to 70% of maximal intake under nonrestricted conditions, increasing pellet cost decreased pellet intake and increased fluid intake more rapidly when the high amphetamine dose was available. Thus, amphetamine was more effective as an economic substitute for pellets when access to pellets was restricted. The response cost for vehicle and both amphetamine concentrations was increased when baboons had nonrestricted and restricted access to pellets. Increasing the response requirement for fluid delivery decreased intake of all three fluids similarly under both pellet-access conditions. The results indicate that substitution between commodities with minimal commonalities can be studied under controlled laboratory conditions and is dependent upon reinforcement schedule and commodity restrictions.     

Food and cocaine self-administration by baboons: effects of alternatives.

The effects of the availability of an alternative reinforcer on responding maintained by food pellets or drug solutions were examined in 8 adult male baboons (Papio hamadrayas anubis). During daily 23-hr experimental sessions, baboons had access to both food pellets and fluid under a two-choice procedure, in which the response requirement, under a fixed-ratio schedule, differed for the two commodities. There were no restrictions on access to water, which was continuously available from a spout at the rear of each cage. In Experiment 1, the fixed-ratio requirement, or cost, for fluid delivery remained constant while the fixed-ratio requirement for pellets was changed every 2 or 3 days when (a) no fluid, (b) a dilute dextrose vehicle, (c) 0.008 mg/kg per delivery cocaine, (d) 0.016 mg/kg per delivery cocaine, or (e) 0.032 mg/kg per delivery cocaine was available concurrently. In Experiment 1, progressively increasing the response requirement for pellets decreased pellet intake, but for 4 baboons pellet intake at maximum pellet cost was lower when cocaine, compared to the vehicle, was available. Increasing the response requirement for pellets had variable effects on vehicle intake. However, increasing the response requirement for pellets increased intake of at least one dose of cocaine to a greater extent than vehicle in all 8 baboons. Thus, cocaine could be considered a more effective economic substitute than vehicle for pellets. Experiment 2 systematically varied the order in which the response requirements for a pellet delivery were presented and added a control condition in which cocaine doses, yoked to the amount self-administered, were given three times during the session by the experimenter. Again, pellet intake at maximal pellet cost was lower when cocaine, compared to the vehicle, was available. In contrast, experimenter-given cocaine doses did not alter responding maintained by pellets. Thus, the effects of self-administered cocaine on responding maintained by food pellets differed from the effects of experimenter-given cocaine on responding maintained by food pellets.     

Stock optimizing: maximizing reinforcers per session on a variable-interval schedule.

In Experiment 1, 2 monkeys earned their daily food ration by pressing a key that delivered food according to a variable-interval 3-min schedule. In Phases 1 and 4, sessions ended after 3 hr. In Phases 2 and 3, sessions ended after a fixed number of responses that reduced food intake and body weights from levels during Phases 1 and 4. Monkeys responded at higher rates and emitted more responses per food delivery when the food earned in a session was reduced. In Experiment 2, monkeys earned their daily food ration by depositing tokens into the response panel. Deposits delivered food according to a variable-interval 3-min schedule. When the token supply was unlimited (Phases 1, 3, and 5), sessions ended after 3 hr. In Phases 2 and 4, sessions ended after 150 tokens were deposited, resulting in a decrease in food intake and body weight. Both monkeys responded at lower rates and emitted fewer responses per food delivery when the food earned in a session was reduced. Experiment 1's results are consistent with a strength account, according to which the phases that reduced body weights increased food's value and therefore increased subjects' response rates. The results of Experiment 2 are consistent with an optimizing strategy, because lowering response rates when food is restricted defends body weight on variable-interval schedules. These contrasting results may be attributed to the discriminability of the contingency between response number and the end of a session being greater in Experiment 2 than in Experiment 1. In consequence, subjects lowered their response rates in order to increase the number of reinforcers per session (stock optimizing).     

Effects of reinforcer magnitude on responding under differential-reinforcement-of-low-rate schedules of rats and pigeons

Experiment I investigated the effects of reinforcer magnitude on differential-reinforcement-of-low-rate (DRL) schedule performance in three phases. In Phase 1, two groups of rats (n = 6 and 5) responded under a DRI. 72-s schedule with reinforcer magnitudes of either 30 or 300 microl of water. After acquisition, the water amounts were reversed for each rat. In Phase 2, the effects of the same reinforcer magnitudes on DRL 18-s schedule performance were examined across conditions. In Phase 3, each rat responded unider a DR1. 18-s schedule in which the water amotnts alternated between 30 and 300 microl daily. Throughout each phase of Experiment 1, the larger reinforcer magnitude resulted in higher response rates and lower reinforcement rates. The peak of the interresponse-time distributions was at a lower value tinder the larger reinforcer magnitude. In Experiment 2, 3 pigeons responded under a DRL 20-s schedule in which reinforcer magnitude (1-s or 6-s access to grain) varied iron session to session. Higher response rates and lower reinforcement rates occurred tinder the longer hopper duration. These results demonstrate that larger reinforcer magnitudes engender less efficient DRL schedule performance in both rats and pigeons, and when reinforcer magnitude was held constant between sessions or was varied daily. The present results are consistent with previous research demonstrating a decrease in efficiency as a function of increased reinforcer magnituide tinder procedures that require a period of time without a specified response. These findings also support the claim that DRI. schedule performance is not governed solely by a timing process.     

Positive conditioned suppression: an explanation in terms of multiple and concurrent schedules

Rats performed under a baseline variable-interval schedule of food presentation. A response-independent food schedule was then superimposed on the baseline schedule for different periods of time across different conditions. The response-independent schedule operated for the whole session in some conditions, intermittently for sixty second periods in some, and intermittently for ten-second periods in others. Under these latter two sets of conditions, the response-independent food schedule was stimulus correlated and alternated with the baseline schedule according to a multiple schedule. Response-independent food presentations always suppressed responding. The degree of suppression tended to increase the longer the period of response-independent food. Control conditions, in which the superimposed schedule was response-dependent, rather than response-independent, did not produce response suppression. The results fit an analysis of positive conditioned suppression phenomena in the context of multiple and concurrent schedule effects.     

Tolerance to the effects of cocaine on performance under behavior-correlated reinforcement magnitude

Four pigeons responded under a fixed-interval 8-min schedule of food delivery in which the amount of food delivered at the end of each interval depended on performance during the interval (i.e., a correlated schedule). Specifically, duration of access to grain was contingent upon the number of responses made during the first 4 min of the interval. This differential outcome did not affect response rates or patterning relative to performance under a simple fixed-interval 8-min schedule. Behavior under the correlated schedule was then investigated under doses of cocaine ranging from 0.3 to 10.0 mg/kg. A bitonic dose-response function was obtained for response rates and the time with head in the food hopper, whereas dose-dependent decreases were observed in the mathematical index of curvature (Fry, Kelleher, & Cook, 1960). The dose that produced the greatest increase in the head-in-hopper time was then administered prior to each session. Following repeated administration of cocaine, disruptions in response patterning were attenuated for all 4 pigeons; tolerance was also observed to the rate-increasing effects and increased head-in-hopper time for 2 pigeons after chronic cocaine administration. Tolerance therefore developed despite the fact that the initial effect of cocaine was to increase the amount of food obtained.     

Bouts of responding on variable-interval schedules: effects of deprivation level

Rats obtained food pellets on a variable-interval schedule of reinforcement by nose poking a lighted key. After training to establish baseline performance (with the mean variable interval set at either 60, 120, or 240 s), the rats were given free access to food during the hour just before their daily session. This satiation operation reduced the rate of key poking. Analysis of the interresponse time distributions (log survivor plots) indicated that key poking occurred in bouts. Prefeeding lengthened the pauses between bouts, shortened the length of bouts (less reliably), and had a relatively small decremental effect on the response rate within bouts. That deprivation level affects mainly between-bout pauses has been reported previously with fixed-ratio schedules. Thus, when the focus is on bouts, the performances maintained by variable-interval schedules and fixed-ratio schedules are similarly affected by deprivation.     

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.     

The effects of differing response types and price manipulations on demand measures

Animals' behavioral needs have become an important component of animal welfare legislation. Behavioral economics provides a framework for the study of such needs. A function, analogous to a demand function relating consumption rate to price, can be obtained by increasing the price (or work) required for access to a commodity. This experiment investigated the effects of different response types and price manipulations on these functions. Six hens pushed a door or pecked a key for food under open economic conditions (short experimental sessions and supplementary food). In Part 1, the number of door pushes required (fixed-ratio schedule) was increased each session, and the force needed to push the door was increased across conditions. In Part 2, the force needed to push the door was increased session to session, and the fixed-ratio schedule was increased across conditions. In Part 3, the number of key pecks required was increased each session. Both response types produced similarly shaped (approximately linear in logarithmic coordinates and downward sloping) demand functions when price was increased by increasing the number of responses required. These imply an elastic demand for food under these conditions. In contrast, increasing the force required to push the door resulted in highly curvilinear functions. These functions indicated little change in consumption across lower door forces and abrupt drops in consumption at higher force requirements, implying mixed elasticity in the animals' demand for food. The differences between the shapes of the two functions seem to arise from the different ways that the two price manipulations alter the time taken to complete the work required. Increasing the fixed-ratio requirement necessarily increases the time needed to complete each response unit, whereas increasing the force requirement does not. The different shapes of the functions were robust when either force or number was varied across sessions and the value of the other was varied over conditions. They were also robust when the price increases were taken from different conditions, showing that the shapes of the functions were independent of the place in the experiment in which the price was examined. Unit price (which combines number and force into a single price measure) unified the data from the two price manipulations to a large degree, producing moderately curved functions. However, there was some variance around the unit price functions, and this was attributable to the different shapes of the underlying functions. The data suggest that different price manipulations may give different measures of animal demand but that unit price might provide some unification.     

A preliminary analysis of adaptive responding under open and closed economies

In the current investigation, we evaluated the effects of open and closed economies on the adaptive behavior of 2 individuals with developmental disabilities. Across both types of economy, progressive-ratio (PR) schedules were used in which the number of responses required to obtain reinforcement increased as the session progressed. In closed-economy sessions, participants were able to obtain reinforcement only through interaction with the PR schedule requirements (i.e., more work resulted in more reinforcer access). In open-economy sessions, participants obtained reinforcers by responding on the PR schedule and were given supplemental (free) access to the reinforcers after completion of the session. In general, more responding was associated with the closed economy.     

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.     

Fixed versus variable sequences of food and stimulus presentation in second-order schedules

Three pigeons were exposed to a second-order schedule in which the behavior specified by a fixed-interval component schedule was reinforced according to a ratio overall schedule. The completion of components not followed by food was signalled by a brief stimulus never paired with food. Food and the stimulus occurred in a random sequence or in fixed alternation, but the overall schedules (variable ratio 2 or fixed ratio 2) ensured that an equal number of food and brief-stimulus presentations occurred in each session. The control exerted by the food and by the brief stimulus was measured by overall response rates, mean pauses, frequency distributions of pauses, and response patterning across components. In general, the stimulus controlled patterns of behavior more similar to those controlled by food when food and the stimulus occurred in a random sequence than when they occurred in fixed alternation.