Rats defend different body weights depending on palatability and accessibility of their food.

Normal adult rats lived on powdered diets adulterated to contain as much as 1.6% quinine sulfate, on a palatable high-fat diet, or in Skinner boxes with 45-mg Noyes pellets available on fixed-ratio (FR) schedules as high as FR 156. They maintained lower body weights over periods of months in proportion to the percentage of quinine adulteration or the fixed ratio. Rats exposed to the high-fat diet overate as much and gained weight as rapidly as rats recovering from food deprivation, and became moderately obese. Rats having become lean or obese contingent on the palatability or accessibility of their diet defended body weight by eating more in the cold, less when force-fed by gavage, and more to restore weight after food deprivation. Yet on chow they restored and defended body weights typical of rats whose diet had been confined to commercially prepared chow. These results are interpreted to be inconsistent with motivational models that rigidly distinguish drive from incentive, that treat body weight changes as evidence for failure to regulate energy balance or body weight, or that rely exclusively on deprivation of food or reduction of body weight for definitions of need for calories. Instead, caloric homeostasis in rats may incorporate ecological constraints.     

Quinine pellets as an inferior good and a Giffen good in rats.

In Experiment 1, 4 rats earned their daily food ration by choosing between two levers. One lever delivered two regular and one quinine-adulterated food pellets, and the other delivered two regular and four quinine pellets. A 20-s intertrial interval separated successive choices. Sessions began with 10 forced trials during which only one lever, selected with p = .5 and cued by a light above it, could deliver its reinforcer. Forced trials were followed by 30 or 150 trials, depending on the condition, during which choices to either lever could be reinforced. Over this range, absolute choice of the four-quinine, two-regular-pellet lever was inversely related to the number of free-choice trials, establishing this reinforcer as an inferior good. In Condition 1 of Experiment 2, the prior design was altered in two ways: (a) one lever delivered four quinine pellets, and the other lever delivered one standard pellet; and (b) sessions ended after 140 free-choice trials. When the number of free-choice trials was reduced to 100 (Condition 2), all 3 rats increased their preference for quinine pellets, confirming their status as an inferior good. In the next several conditions, the number of quinine pellets provided for selecting its associated lever was varied between three and four. Preference for the quinine-pellet alternative was inversely related to the number of pellets it provided, a result defining it as a Giffen good. These findings are not accommodated readily by extant choice models and complicate the search for a unitary model of choice.     

The effect of rearing environments on the contrafreeloading phenomenon in rats

Eight naive rats were reared in enriched or impoverished environments for 39 days after weaning and then lived in operant chambers, in which they could obtain food pellets freely or by lever pressing, for 25 or 30 days. The animals raised in an impoverished environment acquired the bar-press response quickly when placed in the operant chambers and maintained a preference for obtaining food via bar pressing. Animals raised in an enriched environment did not learn to lever press, as demonstrated by low levels of responding and the lack of bar pressing when free food was subsequently removed. It was concluded that restricting animals' postweaning environments facilitated learning in a choice situation, probably because of increased activity levels. The results are interpreted in relation to previous studies on rearing environments and on contrafreeloading.     

The role of the instrumental contingency in the motivational control of performance

In four experiments we investigated an irrelevant incentive effect based upon a transition from hunger to thirst. Hungry rats were trained to lever press either for sucrose solution or for food pellets before performance was tested in extinction while they were thirsty. Reinforcer-specific motivational control was found in the first experiment in that the animals pressed the lever more on tests following training with the sucrose solution rather than with food pellets. Moreover, this effect was seen only when testing was conducted following water, but not following food deprivation. The outcome of the remaining experiments suggests that this motivational control is not mediated by the instrumental contingency between lever pressing and the sucrose reinforcer during training. In these studies lever pressing and chain pulling were reinforced concurrently, one with sucrose and the other with food pellets, in order to equate the noninstrumental functions of the incentives. Following this training, lever pressing in extinction under thirst was unaffected by the type of incentive used as its reinforcer during training.     

Response–reinforcer contiguity versus response-rate–reinforcer-rate covariance in rats' lever pressing: Support for a multiscale view

The multiscale molar view sees behavior as a flow, like a river, extended in time. Matching theory expresses the way activities compete for time. Relative time taken by any activity depends on relative induction. The present experiment tested matching theory applied to concurrent contingent and noncontingent food. As adjunctive activities that compete with operant activity, we recorded hopper head entries and presses on a lever near the food hopper that had no programmed consequences. Eight naïve rats were first exposed to a variable-time 60 s schedule, which across conditions was gradually transformed into a variable-interval 60 s schedule by increasing the proportion of food that was delivered contingent on pressing a lever far from the hopper. Another group of 4 rats that had been trained to press a lever near a food hopper were introduced in the second condition, in which one food delivery was contingent on far-lever pressing. We found induction following a power function to describe pressing on the far lever (operant activity). Matching theory combined with power-function induction also accounted for adjunctive activity. Results with single contingent food deliveries provided little support for the molecular view that behavior consists of discrete responses “strengthened” by immediately following reinforcers.     

Involvement of the rat anterior cingulate cortex in control of instrumental responses guided by reward expectancy

The anterior cingulate cortex (ACC) plays a critical role in stimulus-reinforcement learning and reward-guided selection of actions. Here we conducted a series of experiments to further elucidate the role of the ACC in instrumental behavior involving effort-based decision-making and instrumental learning guided by reward-predictive stimuli. In Experiment 1, rats were trained on a cost-benefit T-maze task in which they could either choose to climb a barrier to obtain a high reward (four pellets) in one arm or a low reward (two pellets) in the other with no barrier present. In line with previous studies, our data reveal that rats with quinolinic acid lesions of the ACC selected the response involving less work and smaller reward. Experiment 2 demonstrates that breaking points of instrumental performance under a progressive ratio schedule were similar in sham-lesioned and ACC-lesioned rats. Thus, lesions of the ACC did not interfere with the effort a rat is willing to expend to obtain a specific reward in this test. In a subsequent task, we examined effort-based decision-making in a lever-press task where rats had the choice between pressing a lever to receive preferred food pellets under a progressive ratio schedule, or free feeding on a less preferred food, i.e. lab chow. Results show that sham- and ACC-lesioned animals had similar breaking points and ingested comparable amounts of less-preferred food. Together, the results of Experiment 1 and 2 suggest that the ACC plays a role in evaluating how much effort to expend for reward; however, the ACC is not necessary in all situations requiring an assessment of costs and benefits. In Experiment 3 we investigated learning and reversal learning of instrumental responses guided by reward predictive stimuli. A reaction time (RT) task demanding conditioned lever release was used in which the upcoming reward magnitude (five vs. one food pellet) was signalled in advance by discriminative visual stimuli. Results revealed that rats with ACC lesions were able to discriminate reward magnitude-predictive stimuli and to adapt instrumental behavior to reversed stimulus-reward magnitude contingencies. Thus, in a simple discrimination task as used here, the ACC appears not to be required to discriminate reward magnitude-predictive stimuli and to use the learned significance of the stimuli to guide instrumental behavior.     

Determinants of reinforcer accumulation during an operant task.

Responses by rats on an earn lever made available food pellets that were delivered to a food cup by responses on a second, collect, lever. The rats could either collect and immediately consume or accumulate (defined as the percentage of multiple earn responses and as the number of pellets earned before a collect response) earned pellets. In Experiment 1, accumulation varied as a function of variations in the earn or collect response requirements and whether the earn and collect levers were proximal (31 cm) or distal (248 cm) to one another. Some accumulation occurred under all but one of the conditions, but generally was higher when the earn and collect levers were distal to one another, particularly when the earn response requirement was fixed-ratio (FR) 1. In Experiment 2, the contributions of responses and time to accumulation were assessed by comparing an FR 20 earn response requirement to a condition in which only a single earn response was required at the end of a time interval nominally yoked to the FR interval. When 248 cm separated the earn and collect levers, accumulation was always greater in the FR condition, and it was not systematically related to reinforcement rate. In Experiment 3, increasing the earn response requirement with a progressive-ratio schedule that reset only with a collect response increased the likelihood of accumulation when the collect and earn levers were 248 cm apart, even though such accumulation increased the next earn response requirement. Reinforcer accumulation is an understudied dimension of operant behavior that relates to the analysis of such phenomena as hoarding and self-control, in that they too involve accumulating versus immediately collecting or consuming reinforcers.     

Effects of haloperidol on schedule-induced polydipsia.

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

Diversity and substitutability of adjunctive activities under fixed-interval schedules of food reinforcement

Six rats received food contingent on pressing a lever on fixed-ratio 1, fixed-interval 30-second, and fixed-interval 60-second schedules, with concurrent access to a drinking spout, a running wheel, and a block of wood. Drinking, running, and chewing were monitored automatically, and these and other activities were observed directly during selected sessions. Because all sessions ended after delivery of 60 pellets, total time available for activities other than eating increased over the three schedules. Time spent contacting the lever and visiting the food tray increased in proportion to total available time, whereas the time spent in other activities changed in a complex manner such that drinking was the dominant adjunctive behavior in the 30-second condition, and running or chewing the dominant adjunctive behavior in five of six rats in the 60-second condition. General activity and grooming also occupied significant amounts of time. In a subsequent part of the experiment, running and chewing were prevented, and the majority of other activities, especially drinking and grooming, increased. The results show that (a) FI schedules of food reinforcement are accompanied by a wide variety of adjunctive activities; (b) the preferred activity differs according to the schedule duration; and (c) the extent to which activities substitute for one another is limited by the tendency for different activities to occupy different parts of the interreinforcement interval.     

Alcohol seeking by rats: Action or habit?

In two experiments, we examined the relative susceptibility to outcome devaluation of lever pressing by rats for either a 10% ethanol solution or food pellets. The rats were trained to press different levers for these two reinforcers using a sucrose-substitution procedure. An aversion was then conditioned from either the ethanol solution or the food pellets by pairing consumption with illness induced by lithium chloride. When instrumental performance was subsequently tested in extinction, the rats pressed less on the pellet lever if the pellets, rather than the ethanol, had been devalued by aversion conditioning. By contrast, performance on the ethanol lever was unaffected by whether the ethanol or pellets were devalued. Moreover, noncontingent presentations of the devalued reinforcer had no impact on test performance. The differential resistance to outcome devaluation suggests that, in contrast to food seeking, alcohol seeking is a stimulus-response habit rather than a goal-directed action mediated by a representation of the action-outcome contingency.     

Incentive learning and the motivational control of instrumental performance

Hungry rats were trained to press a lever and pull a chain concurrently, with one action being reinforced with a sucrose solution and the other with food pellets. In addition, in the first two experiments all animals experienced non-contingent presentations of the two incentives in the absence of the operant manipulanda while either thirsty or hungry and either before (Experiment 1A) or after (Experiment 1B) the instrumental training. When lever pressing was assessed subsequently in extinction under thirst, the animals pressed at a relatively high rate only if (1) this action had been reinforced with the sucrose solution rather than the food pellets during training and (2) they had received the non-contingent presentations of the sucrose solution and food pellets on days on which they were thirsty rather than hungry. A third experiment demonstrated that non-contingent exposure to the sucrose solution alone, but not to water under thirst was sufficient to bring about this type of motivational control of instrumental performance.     

The generality of selective observing

Four rats obtained food pellets by poking a key and 5-s presentations of the discriminative stimuli by pressing a lever. Every 1 or 2 min, the prevailing schedule of reinforcement for key poking alternated between rich (either variable-interval [VI] 30 s or VI 60 s) and lean (either VI 240 s, VI 480 s, or extinction) components. While the key was dark (mixed-schedule stimulus), no exteroceptive stimulus indicated the prevailing schedule. A lever press (i.e., an observing response), however, illuminated the key for 5 s with either a steady light (S+), signaling the rich reinforcement schedule, or a blinking light (S-), signaling the lean reinforcement schedule. One goal was to determine whether rats would engage in selective observing (i.e., a pattern of responding that maintains contact with S+ and decreases contact with S-). Such a pattern was found, in that a 5-s presentation of S+ was followed relatively quickly by another observing response (which likely produced another 5-s period of S+), whereas exposure to S- resulted in extended breaks from observing. Additional conditions demonstrated that the rate of observing remained high when lever presses were effective only when the rich reinforcement schedule was in effect (S+ only condition), but decreased to a low level when lever presses were effective only during the lean reinforcement component (S- only condition) or when lever presses had no effect (in removing the mixed stimulus or presenting the multiple-schedule stimuli). These findings are consistent with relativistic conceptualizations of conditioned reinforcement and extend the generality of selective observing to procedures in which the experimenter controls the duration of stimulus presentations, the schedule components both offer intermittent food reinforcement, and rats serve as subjects.     

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.     

Effects of response-dependent and independent electric shock on schedule-induced polydipsia

In Experiment I, lever pressing by rats was maintained by the delivery of food pellets under a 45-sec fixed-interval schedule. Fixed-time 180-sec and fixed-interval 180-sec schedules of shock delivery were systematically superimposed on the baseline food schedule to study effects on schedule-induced water intake. Response-dependent shock had little, if any, effect on water intake, whereas shocks independent of lever pressing attenuated fluid intake. In Experiment 2, rats received food pellets under a fixed-time 60-sec schedule. Electric shock delivered concurrently under a variable-time 180-sec schedule, but never while the animal was licking or within 5 sec after licking terminated, led to similar attenuation of water intake. These findings suggest that schedule-induced polydipsia is sensitive to differences in the functional properties of response-independent and dependent electric shock.     

Thiamin deprivation in ventromedial hypothalamic hyperphagic rats: anorexia, specificity of food aversion, and a dietary consideration.

The anorexic consequence of thiamin deprivation was investigated in ventromedial hypothalamic (VMH) hyperphagic rats under either high-fat or low-fat thiamin-free diet conditions. The low-fat diet maintained feeding significantly longer in thiamin-deprived VMH rats than in intact rats, whereas the hig-fat diet sustained feeding in thiamin-deficient intact rats and accelerated anorexia onset in vitamin B1 deprived VMH rats. This effect was noted under both ad lib and pair-feeding conditions. Thiamin-deprived VMH rats subjected to weight control developed anorexia sooner than intact subjects regardless of the diet employed. The VMH rats fed a high-fat diet failed to resume feeding after thiamin readministration, which was interpreted as a permanent aversion to this diet. The relation between dietary intake and conditioned taste aversion is discussed with reference to the VMH and intact rat.     

A comparison of operant licking and lever pressing in the rat

Two groups of rats were trained on a signaled, free-operant, avoidance procedure to lick or to lever press in order to avoid shock while water-deprived or satiated and, in the case of licking, while ingesting deionized water, isotonic saline, or 10% sucrose. The most effective avoidance licking occurred while the rats were water-deprived and ingesting 10% sucrose. Water deprivation level had no effect on lever pressing for shock avoidance. Two other groups of rats were operantly conditioned to lick or to lever press for food pellets while waterdeprived or satiated and, in the case of licking, while ingesting deionized water or 10% sucrose. The most effective licking for food reinforcement occurred while the rats were water-deprived and ingesting 10% sucrose. Water deprivation level had no effect on lever pressing for food reinforcement. The data indicated that effective operant licking must be supported by factors related to water regulation and taste palatability.     

Is induction produced by upcoming food-pellet reinforcement the outcome of an increase in overall activity or in operant responding?

Researchers have demonstrated that rats reliably increase their rates of pressing a lever for 1% liquid-sucrose reinforcement if they will soon have the opportunity to press a lever for food-pellet reinforcement. In the present experiments, the authors investigated if this increase in response rates occurred because the upcoming food pellets produced an increase in all behaviors (i.e., general arousal) or an increase in only the specific operant response (i.e., lever pressing). The results of Experiments 1 and 2 showed that the appearance of induction in rats' lever pressing for 1% sucrose reinforcement when food-pellet reinforcement was upcoming did not coincide with increases in the frequency of running in a wheel or making a nonreinforced nose-poke response. On the other hand, in Experiment 3, the authors found the appearance of induction coincided with increase nonreinforced lever presses on an adjacent lever. These results shed doubt on the idea that induction is a result of a general increase in all activity, and suggest instead that the increase in responding that occurs during induction is limited to the operant response.     

Overshadowing of a context aversion by a novel incentive in operant conditioning

Two experiments examined the processes underlying the suppression of instrumental behaviours by lithium in rats, as reported by Meachum (1988 and this issue). Experiment 1 examined whether presenting a novel sucrose solution prior to lithium chloride administration would overshadow aversion learning to either the stimuli of the operant chamber or to familiar food pellets. After lever pressing had been established, and in the absence of responding, animals received free deliveries of a novel sucrose solution, familiar food pellets, or both, or they were exposed to only the cues of the operant chamber, prior to lithium injections. Lever pressing for food pellets was then assessed. It was found that the animals receiving the novel sucrose, either alone or with the familiar food pellets, pressed more for pellets than either the group receiving only food pellets or the group exposed to only the context. In addition, there was no appreciable difference in the response rates between the context-only group and the group that received the familiar food pellets. These outcomes were interpreted in terms of the novel sucrose overshadowing aversion learning to the context. Experiment 2 investigated whether in fact aversive contextual conditioning could be obtained using the present parameters. This was accomplished by directly manipulating the contexts. In this experiment animals were trained to lever press in two distinctive contexts. Subsequently, one context was paired with the novel sucrose, and the second was experienced in the absence of reinforcement prior to toxicosis. During a subsequent non-reinforced test it was found that responding in the context paired with the novel sucrose was considerably higher than responding in the context that was experienced alone. These findings stand in contrast to the taste-mediated contextual potentiation observed when a consumatory response is used to assess aversive contextual conditioning.     

Spaced food but not electrical brain stimulation induces polydipsia and air licking.

An attempt was made to induce polydipsia in rats whose lever pressing was reinforced with food pellets or electrical brain stimulation. Nine food-deprived, water-sated rats drank water excessively during sessions in which food pellets were delivered. When brain stimulation was substituted for food, drinking immediately ceased. Delivering brain stimulation according to a variety of schedules, pairing brain stimulation with food reinforcement, and substituting an air stream for water, each failed to produce polydipsic licking. These results show that polydipsia is not induced by all reinforcers.     

Effect of ventromedial hypothalamic procaine injections on feeding, lever pressing, and other behavior in rats.

Rats were given bilateral injections of 1-2 mul procaine HCl solution (50 mug/mul) in the region of the ventromedial hypothalamus. Normal-sized meals occurred reliably with a latency of 16-120 sec, indicating that this region has the function of inhibiting onset of eating, not just the function of stopping a meal. Food-rewarded fixed-ratio-of-one (FR 1; continuous reinforcement) lever pressing was elicited by 2-mul but not by 1-mul procaine. The FR 64 pressing appeared to be disrupted during a period of increased activity following procaine injection, although visual observations indicated that the pressing which did occur was normal in topography and was significantly associated in temporal sequence with approaches of the food magazine.