Effects of acute and chronic flunitrazepam on delay discounting in pigeons

Delay to delivery of a reinforcer can decrease responding for that reinforcer and increase responding for smaller reinforcers that are available concurrently and delivered without delay; acute administration of drugs can alter responding for large, delayed reinforcers, although the impact of chronic treatment on delay discounting is not well understood. In this experiment, the effects of repeated administration of the benzodiazepine flunitrazepam were studied in 6 pigeons responding on one key to receive food that was delivered immediately and on a second key to receive a larger amount of food that was delivered following delays which increased across a single session. Pigeons responded predominantly for the large reinforcer when there were no delays and when delays were short; however, as delays increased, responding for the large reinforcer decreased. Acutely, flunitrazepam (0.32, 1.0 and 3.2 mg/kg) dose-dependently increased responding for the large reinforcer, shifting the discounting curve rightward and upward. Repeated administration of flunitrazepam (0.32, 1.0 and 3.2 mg/kg, each for six sessions, separated by one session during which vehicle was administered) did not markedly alter its effects on responding for the large reinforcer, indicating that the development of tolerance to this effect of flunitrazepam is modest under these conditions.     

Effects of reinforcer probability, delay, and response requirements on the choices of rats and pigeons: possible species differences

In Experiment 1 with rats, a left lever press led to a 5-s delay and then a possible reinforcer. A right lever press led to an adjusting delay and then a certain reinforcer. This delay was adjusted over trials to estimate an indifference point, or a delay at which the two alternatives were chosen about equally often. Indifference points increased as the probability of reinforcement for the left lever decreased. In some conditions with a 20% chance of food, a light above the left lever was lit during the 5-s delay on all trials, but in other conditions, the light was only lit on those trials that ended with food. Unlike previous results with pigeons, the presence or absence of the delay light on no-food trials had no effect on the rats' indifference points. In other conditions, the rats showed less preference for the 20% alternative when the time between trials was longer. In Experiment 2 with rats, fixed-interval schedules were used instead of simple delays, and the presence or absence of the fixed-interval requirement on no-food trials had no effect on the indifference points. In Experiment 3 with rats and Experiment 4 with pigeons, the animals chose between a fixed-ratio 8 schedule that led to food on 33% of the trials and an adjusting-ratio schedule with food on 100% of the trials. Surprisingly, the rats showed less preference for the 33% alternative in conditions in which the ratio requirement was omitted on no-food trials. For the pigeons, the presence or absence of the ratio requirement on no-food trials had little effect. The results suggest that there may be differences between rats and pigeons in how they respond in choice situations involving delayed and probabilistic reinforcers.     

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.     

Choice with delayed and probabilistic reinforcers: effects of prereinforcer and postreinforcer stimuli.

In Experiment 1, pigeons' pecks on a green key led to a 5-s delay with green houselights, and then food was delivered on 20% (or, in other conditions, 50%) of the trials. Pecks on a red key led to an adjusting delay with red houselights, and then food was delivered on every trial. The adjusting delay was used to estimate indifference points: delays at which the two alternatives were chosen about equally often. Varying the presence or absence of green houselights during the delays that preceded possible food deliveries had large effects on choice. In contrast, varying the presence of the green or red houselights in the intertrial intervals had no effects on choice. In Experiment 2, pecks on the green key led to delays of either 5 s or 30 s with green houselights, and then food was delivered on 20% of the trials. Varying the duration of the green houselights on nonreinforced trials had no effect on choice. The results suggest that the green houselights served as a conditioned reinforcer at some times but not at others, depending on whether or not there was a possibility that a primary reinforcer might be delivered. Given this interpretation of what constitutes a conditioned reinforcer, most of the results were consistent with the view that the strength of a conditioned reinforcer is inversely related to its duration.     

Overshadowing of a stimulus-reinforcer association by an instrumental response

In two experiments, rats were first exposed to pairings of a clicker and food; they were subsequently, in order to measure the effectiveness of the clicker as a conditioned reinforcer, given the opportunity to press a lever which turned the clicker on. For one group of animals the food originally delivered in the presence of the clicker had been contingent on their performance of an instrumental response (running in a running wheel); for a second the contingency between clicker and food had been purely classical. Although the actual correlation between clicker and food was identical for the two groups, the clicker was a less effective conditioned reinforcer for the first group than for the second. In a third experiment, all animals were initially required to run to obtain food in the presence of the clicker, but one group received additional trials on which food was delivered contingent on running in the absence of the clicker. This group showed less tendency to lever press for the clicker than a second group that had received free food on trials when the clicker was not presented. The results of all three experiments suggest that conditioning to the clicker could be overshadowed if the occurrence of food was more reliably predicted by the execution of an instrumental running response; they thus support the view that instrumental conditioning depends on the establishment of an association between response and reinforcer similar to the association between stimulus and reinforcer underlying classical conditioning.     

Theories of probabilistic reinforcement.

In three experiments, pigeons chose between two alternatives that differed in the probability of reinforcement and the delay to reinforcement. A peck at a red key led to a delay of 5 s and then a possible reinforcer. A peck at a green key led to an adjusting delay and then a certain reinforcer. This delay was adjusted over trials so as to estimate an indifference point, or a duration at which the two alternatives were chosen about equally often. In Experiments 1 and 2, the intertrial interval was varied across conditions, and these variations had no systematic effects on choice. In Experiment 3, the stimuli that followed a choice of the red key differed across conditions. In some conditions, a red houselight was presented for 5 s after each choice of the red key. In other conditions, the red houselight was present on reinforced trials but not on nonreinforced trials. Subjects exhibited greater preference for the red key in the latter case. The results were used to evaluate four different theories of probabilistic reinforcement. The results were most consistent with the view that the value or effectiveness of a probabilistic reinforcer is determined by the total time per reinforcer spent in the presence of stimuli associated with the probabilistic alternative. According to this view, probabilistic reinforcers are analogous to reinforcers that are delivered after variable delays.     

Effects of methylphenidate and morphine on delay-discount functions obtained within sessions

Four rats responded under a "self-control" procedure designed to obtain delay-discount functions within sessions. Each session consisted of seven blocks, with seven trials within each block. Each block consisted of two initial forced-choice trials followed by five free-choice trials. On choice trials, the rats could press either of two retractable levers. A press on one lever was followed by presentation of a smaller reinforcer (a single dipper presentation of a sucrose solution); a press on the other lever was followed by presentation of a larger reinforcer (four consecutive dipper presentations). The delay associated with the smaller reinforcer always was 0 s, whereas the signaled delay associated with the larger reinforcer increased across blocks (from 0 to 50 s). Under these conditions, the percentage of choices of the larger reinforcer decreased across blocks, and relatively reliable delay-discount functions were obtained within sessions. Doses of methylphenidate (1.0 to 17.0 mg/kg) and morphine (0.3 to 17.0 mg/kg) were then administered prior to selected sessions. Typically, intermediate doses of methylphenidate shifted the discount functions to the right (increased choices of the larger reinforcer). For 2 of the rats, this effect was pronounced; for the other 2 rats, this effect occurred after the range of delays for the larger reinforcer was decreased (0 to 20 s). On the other hand, in most cases morphine produced a slight leftward shift in the discount function (decreased choices of the larger reinforcer). The present procedure appears to be a useful and efficient method to characterize drug effects on an entire delay-discount function. As with many procedures used to study self-control choices, however, sources of control other than reinforcement delay and amount may have been operating in the present study, and these sources must be considered when interpreting drug effects.     

Behavioral and pharmacological variables affecting risky choice in rats.

The effects of manipulations of response requirement, intertrial interval (ITI), and psychoactive drugs (ethanol, phencyclidine, and d-amphetamine) on lever choice under concurrent fixed-ratio schedules were investigated in rats. Responding on the "certain' lever produced three 45-mg pellets, whereas responding on the "risky" lever produced either 15 pellets (p = .33) or no pellets (p .67). Rats earned all food during the session, which ended after 12 forced trials and 93 choice trials or 90 min, whichever occurred first. When the response requirement was increased from 1 to 16 and the ITI was 20 s, percentage of risky choice was inversely related to fixed-ratio value. When only a single response was required but the ITI was manipulated between 20 and 120 s (with maximum session duration held constant), percentage of risky choice was directly related to length of the ITI. The effects of the drugs were investigated first at an ITI of 20 s, when risky choice was low for most rats, and then at an ITI of 80 s, when risky choice was higher for most rats. Ethanol usually decreased risky choice. Phencyclidine did not usually affect risky choice when the ITI was 20 s but decreased it in half the rats when the ITI was 80 s. For d-amphetamine, the effects appeared to he related to baseline probability of risky choice; that is, low probabilities were increased and high probabilities were decreased. Although increase in risky choice as a function of the ITI is at variance with previous ITI data, it is consistent with foraging data showing that risk aversion decreases as food availability decreases. The pharmacological manipulations showed that drug effects on risky choice may be influenced by the baseline probability of risky choice, just as drug effects can be a function of baseline response rate.     

ISOLATING BEHAVIORAL MECHANISMS OF INTERTEMPORAL CHOICE: NICOTINE EFFECT ON DELAY DISCOUNTING AND AMOUNT SENSITIVITY

Many drugs of abuse produce changes in impulsive choice, that is, choice for a smaller—sooner reinforcer over a larger—later reinforcer. Because the alternatives differ in both delay and amount, it is not clear whether these drug effects are due to the differences in reinforcer delay or amount. To isolate the effects of delay, we used a titrating delay procedure. In phase 1, 9 rats made discrete choices between variable delays (1 or 19 s, equal probability of each) and a delay to a single food pellet. The computer titrated the delay to a single food pellet until the rats were indifferent between the two options. This indifference delay was used as the starting value for the titrating delay for all future sessions. We next evaluated the acute effects of nicotine (subcutaneous 1.0, 0.3, 0.1, and 0.03 mg/kg) on choice. If nicotine increases delay discounting, it should have increased preference for the variable delay. Instead, nicotine had very little effect on choice. In a second phase, the titrated delay alternative produced three food pellets instead of one, which was again produced by the variable delay (1 s or 19 s) alternative. Under this procedure, nicotine increased preference for the one pellet alternative. Nicotine‐induced changes in impulsive choice are therefore likely due to differences in reinforcer amount rather than differences in reinforcer delay. In addition, it may be necessary to include an amount sensitivity parameter in any mathematical model of choice when the alternatives differ in reinforcer amount.     

Within-session changes in key and lever pressing for water during several multiple variable-interval schedules

Rats pressed keys or levers for water reinforcers delivered by several multiple variable-interval schedules. The programmed rate of reinforcement varied from 15 to 240 reinforcers per hour in different conditions. Responding usually increased and then decreased within experimental sessions. As for food reinforcers, the within-session changes in both lever and key pressing were smaller, peaked later, and were more symmetrical around the middle of the session for lower than for higher rates of reinforcement. When schedules provided high rates of reinforcement, some quantitative differences appeared in the within-session changes for lever and key pressing and for food and water. These results imply that basically similar factors produce within-session changes in responding for lever and key pressing and for food and water. The nature of the reinforcer and the choice of response can also influence the quantitative properties of within-session changes at high rates of reinforcement. Finally, the results show that the application of Herrnstein's (1970) equation to rates of responding averaged over the session requires careful consideration.     

Conditioned reinforcement and choice

In a series of three experiments, rats were exposed to successive schedule components arranged on two levers, in which lever pressing produced a light, and nose-key pressing produced water in 50% of the light periods. When one auditory signal was presented only during those light periods correlated with water on one lever, and a different signal was presented only during those light periods correlated with nonreinforcement on the other lever, the former lever was preferred in choice trials, and higher rates of responding were maintained on the former lever in nonchoice (forced) trials. Thus, the rats preferred a schedule component that included a conditioned reinforcer over one that did not, with the schedules of primary reinforcement and the information value of the signals equated. Preferences were maintained when one or the other of the auditory signals was deleted, but were not established in naive subjects when training began with either the positive or negative signal only. Discriminative control of nose-key pressing by the auditory signals was highly variable across subjects and was not correlated with choice.     

Determinants of pigeons' choices in token-based self-control procedures

Four pigeons were exposed to a token-based self-control procedure with stimulus lights serving as token reinforcers. Smaller-reinforcer choices produced one token immediately; larger-reinforcer choices produced three tokens following a delay. Each token could be exchanged for 2-s access to food during a signaled exchange period each trial. The main variables of interest were the exchange delays (delays from the choice to the exchange stimulus) and the food delays (also timed from the choice), which were varied separately and together across blocks of sessions. When exchange delays and food delays were shorter following smaller-reinforcer choices, strong preference for the smaller reinforcer was observed. When exchange delays and food delays were equal for both options, strong preference for the larger reinforcer was observed. When food delays were equal for both options but exchange delays were shorter for smaller-reinforcer choices, preference for the larger reinforcer generally was less extreme than under conditions in which both exchange and food delays were equal. When exchange delays were equal for both options but food delays were shorter for smaller-reinforcer choices, preference for the smaller reinforcer generally was less extreme than under conditions in which both exchange and food delays favored smaller-reinforcer choices. On the whole, the results were consistent with prior research on token-based self-control procedures in showing that choices are governed by reinforcer immediacy when exchange and food delays are unequal and by reinforcer amount when exchange and food delays are equal. Further, by decoupling the exchange delays from food delays, the results tentatively support a role for the exchange stimulus as a conditioned reinforcer.     

"Turning back the clock" on serial-stimulus sign tracking.

Two experiments examined the effects of a negative (setback) response contingency on key pecking engendered by a changing light-intensity stimulus clock (ramp stimulus) signaling fixed-time 30-s food deliveries. The response contingency specified that responses would immediately decrease the light-intensity value, and, because food was delivered only after the highest intensity value was presented, would delay food delivery by 1 s for each response. The first experiment examined the acquisition and maintenance of responding for a group trained with the contingency in effect and for a group trained on a response-independent schedule with the ramp stimulus prior to introduction of the contingency. The first group acquired low rates of key pecking, and, after considerable exposure to the contingency, those rates were reduced to low levels. The rates of responding for the second group were reduced very rapidly (within four to five trials) after introduction of the setback contingency. For both groups, rates of responding increased for all but 1 bird when the contingency was removed. A second experiment compared the separate effects of each part of the response contingency. One group was exposed only to the stimulus setback (stimulus only), and a second group was exposed only to the delay of the reinforcer (delay only). The stimulus-only group's rates of responding were immediately reduced to moderate levels, but for most of the birds, these rates recovered quickly when the contingency was removed. The delay-only groups's rates decreased after several trials, to very low levels, and recovery of responding took several sessions once the contingency was removed. The results suggest that (a) sign-tracking behavior elicited by an added clock stimulus may be reduced rapidly and persistently when a setback contingency is imposed, and (b) the success of the contingency is due both to response-dependent stimulus change and response-dependent alterations in the frequency of food delivery. The operation of the contingency is compared with the effects of secondary reinforcement and punishment procedures.     

Procrastination by pigeons: preference for larger, more delayed work requirements.

In three experiments, pigeons chose between alternatives that required the completion of a small ratio schedule early in the trial or a larger ratio schedule later in the trial. Completion of the ratio requirement did not lead to an immediate reinforcer, but simply allowed the events of the trial to continue. In Experiment 1, the ratio requirements interrupted periods in which food was delivered on a variable-time schedule. In Experiments 2 and 3, each ratio requirement was preceded and followed by a delay, and only one reinforcer was delivered, at the end of each trial. Two of the experiments used an adjusting-ratio procedure in which the ratio requirement was increased and decreased over trials so as to estimate an indifference point--a ratio size at which the two alternatives were chosen about equally often. These experiments found clear evidence for "procrastination"--the choice of a larger but more delayed response requirement. In some cases, subjects chose the more delayed ratio schedule even when it was larger than the more immediate alternative by a factor of four or more. The results suggest that as the delay to the start of a ratio requirement is increased, it has progressively less effect on choice behavior, in much the same way that delaying a positive reinforcer reduces it effect on choice.     

A choice procedure to assess the aversive effects of drugs in rodents

The goal of this series of experiments was to develop an operant choice procedure to examine rapidly the punishing effects of intravenous drugs in rats. First, the cardiovascular effects of experimenter‐administered intravenous histamine, a known aversive drug, were assessed to determine a biologically active dose range. Next, rats responded on each of two levers with concurrently available fixed‐ratio 1 schedules of food reinforcement. Intravenous histamine was delivered along with food when responses were made on one of the options, and the lever on which both food and histamine were contingent was switched on a regular basis. A dose of 1.0 mg/kg/inj of histamine was effective in moving responding to the alternate lever, whereas saline, 0.1, or 0.3 mg/kg/inj of histamine were not. Histamine injections produced reliable selection of the alternate lever when they were presented on the same lever for three consecutive sessions, but not when they were switched between levers on each session. In addition, histamine produced greater selection of the alternate lever when it was presented with shorter intertrial interval durations. These findings indicate that, with appropriate parameters, the aversive effects of histamine and perhaps other drugs can be established rapidly using a concurrent choice procedure.     

Response duration is sensitive to both immediate and delayed reinforcement

We investigated the duration of lever pressing by rats when the delivery of appetitive reinforcers was contingent upon response duration. In the first experiment, response durations increased when duration requirements were imposed, and they decreased when duration requirements were removed. This effect occurred whether reinforcers were immediate or delayed by 8 s. In order to maintain the integrity of the delay intervals, reinforcer delivery was dependent upon both lever depression and release. In a second experiment, lever depression only and a response duration of at least 4 s were required for reinforcer delivery. Compared to immediate reinforcement conditions, delayed reinforcers increased both variability and the length of the maximum response durations. In a third experiment, immediate reinforcers were delivered contingent upon lever depression and release under a variety of duration requirements. Median lever‐press durations tracked the contingencies rapidly. Across all three experiments, rats emitted numerous response durations that were too short to satisfy the reinforcer requirements, and bimodal distributions similar to those produced by differential reinforcement of low rate schedules were evident for most rats. In many aspects, response duration responds to reinforcement parameters in a fashion similar to rate of discrete responding, but an examination of this continuous dimension of behavior may provide additional information about environment–behavior relationships.     

Self-control in mentally retarded adolescents: choice as a function of amount and delay of reinforcement.

Three severely mentally retarded adolescents were studied under discrete-trial procedures in which a choice was arranged between edible reinforcers that differed in magnitude and, in some conditions, delay. In the absence of delays the larger reinforcer was consistently chosen. Under conditions in which the smaller reinforcer was not delayed, increasing the delay to delivery of the larger reinforcer decreased the percentage of trials in which that reinforcer was chosen. All subjects directed the majority of choice responses to the smaller reinforcer when the larger reinforcer was sufficiently delayed, although the value at which this occurred differed across subjects. Under conditions in which the larger reinforcer initially was sufficiently delayed to result in preference for the smaller one, progressively increasing in 5-s increments the delay to both reinforcers increased percentage of trials with the larger reinforcer chosen. At sufficiently long delays, 2 of the subjects consistently chose the larger, but more delayed, reinforcer, and the 3rd subject chose that reinforcer on half of the trials. These results are consistent with the findings of prior studies in which adult humans responded to terminate noise and pigeons responded to produce food.     

Fix and sample with rats in the dynamics of choice

The generality of the molar view of behavior was extended to the study of choice with rats, showing the usefulness of studying order at various levels of extendedness. Rats' presses on two levers produced food according to concurrent variable-interval variable-interval schedules. Seven different reinforcer ratios were arranged within each session, without cues identifying them, and separated by blackouts. To alternate between levers, rats pressed on a third changeover lever. Choice changed rapidly with changes in component reinforcer ratio, and more presses occurred on the lever with the higher reinforcer rate. With continuing reinforcers, choice shifted progressively in the direction of the reinforced lever, but shifted more slowly with each new reinforcer. Sensitivity to reinforcer ratio, as estimated by the generalized matching law, reached an average of 0.9 and exceeded that documented in previous studies with pigeons. Visits to the more-reinforced lever preceded by a reinforcer from that lever increased in duration, while all visits to the less-reinforced lever decreased in duration. Thus, the rats' performances moved faster toward fix and sample than did pigeons' performances in previous studies. Analysis of the effects of sequences of reinforcer sources indicated that sequences of five to seven reinforcers might have sufficed for studying local effects of reinforcers with rats. This study supports the idea that reinforcer sequences control choice between reinforcers, pulses in preference, and visits following reinforcers.     

The effects of constant versus varied reinforcers on preference and resistance to change

Previous research has demonstrated that factors such as reinforcer frequency, amount, and delay have similar effects on resistance to change and preference. In the present study, 4 boys with autism made choices between a constant reinforcer (one that was the same food item every trial) and a varied food reinforcer (one that varied randomly between three possible food items). For all 4 boys, varied reinforcers were preferred over constant reinforcers, and they maintained higher response rates than constant reinforcers. In addition, when a distraction (a video clip) was introduced, responding maintained by varied reinforcers was more resistant to distraction than responding maintained by constant reinforcers. Thus, the present experiment extended the generality of the relation between preference and resistance to change to variation in reinforcer quality.