Delay discounting in Lewis and Fischer 344 rats: steady-state and rapid-determination adjusting-amount procedures

Lewis rats have been shown to make more impulsive choices than Fischer 344 rats in discrete-trial choice procedures that arrange fixed (i.e., nontitrating) reinforcement parameters. However, nontitrating procedures yield only gross estimates of preference, as choice measures in animal subjects are rarely graded at the level of the individual subject. The present study was designed to examine potential strain differences in delay discounting using an adjusting-amount procedure, in which distributed (rather than exclusive) choice is observed due to dynamic titration of reinforcer magnitude across trials. Using a steady-state version of the adjusting-amount procedure in which delay was manipulated between experimental conditions, steeper delay discounting was observed in Lewis rats compared to Fischer 344 rats; further, delay discounting in both strains was well described by the traditional hyperbolic discounting model. However, upon partial completion of the present study, a study published elsewhere (Wilhelm & Mitchell, 2009) demonstrated no difference in delay discounting between these strains with the use of a more rapid version of the adjusting-amount procedure (i.e., in which delay is manipulated daily). Thus, following completion of the steady-state assessment in the present study, all surviving Lewis and Fischer 344 rats completed an approximation of this rapid-determination procedure in which no strain difference in delay discounting was observed.     

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.     

Responding during reinforcement delay in a self-control paradigm.

Eight pigeons chose between a small, immediate reinforcer and a large, increasingly delayed reinforcer. Responding during the large-reinforcer delays was examined. During large-reinforcer delays, pecks on one key produced the small, immediate reinforcer; pecks on the other key had no effect. Thus, a pigeon could reverse its initial choice of the large, delayed reinforcer, or it could maintain its original choice. Pigeons that made a relatively high number of initial large-reinforcer choices tended to maintain these choices, and those pigeons that actually received a relatively high number of large reinforcers, tended to respond more frequently on the ineffective key during the delay periods. The findings suggest that some previous studies of self-control training in pigeons may have resulted in increased self-control partially due to a lack of opportunity for the pigeons to change their choices.     

Effects of pre-trial response requirements on self-control choices by rats and pigeons

Parallel experiments with rats and pigeons examined whether the size of a pre-trial ratio requirement would affect choices in a self-control situation. In different conditions, either 1 response or 40 responses were required before each trial. In the first half of each experiment, an adjusting-ratio schedule was used, in which subjects could choose a fixed-ratio schedule leading to a small reinforcer, or an adjusting-ratio schedule leading to a larger reinforcer. The size of the adjusting ratio requirement was increased and decreased over trials based on the subject's responses, in order to estimate an indifference point-a ratio at which the two alternatives were chosen about equally often. The second half of each experiment used an adjusting-delay procedure-fixed and adjusting delays to the small and large reinforcers were used instead of ratio requirements. In some conditions, particularly with the reinforcer delays, the rats had consistently longer adjusting delays with the larger pre-trial ratios, reflecting a greater tendency to choose the larger, delayed reinforcer when more responding was required to reach the choice point. No consistent effects of the pre-trial ratio were found for the pigeons in any of the conditions. These results may indicate that rats are more sensitive to the long-term reinforcement rates of the two alternatives, or they may result from a shallower temporal discounting rate for rats than for pigeons, a difference that has been observed in previous studies.     

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.     

Transitional and steady-state choice behavior under an adjusting-delay schedule

Twelve rats made repeated choices on an adjusting-delay schedule between a smaller reinforcer (A) that was delivered immediately after a response and a larger reinforcer (B) that was delivered after a delay which increased or decreased by 20% depending on the subject's choices in successive blocks of trials. In two phases of the experiment (100 sessions and 40 sessions), reinforcer sizes were selected which enabled theoretical parameters expressing the rate of delay discounting and sensitivity to reinforcer size to be estimated from the ratio of the indifference delays obtained in the two phases. Indifference delays, calculated from adjusting delays in the last 10 sessions of each phase, were shorter when the sizes of A and B were 14 and 25 μl of a 0.6 M sucrose solution than when they were 25 and 100 μl of the same solution. The ratio of the indifference delays was significantly smaller than that predicted on the basis of an assumed linear relation between reinforcer size and instantaneous reinforcer value, consistent with a previous proposal that this relation may be hyperbolic in form. Estimates of the rate of delay discounting based on the ratio of the two indifference delays (mean, 0.08 s(-1)) were similar to values obtained previously using different intertemporal choice protocols. Estimates of the size-sensitivity parameter (mean 113 μl) were similar to estimates recently derived from performance on progressive-ratio schedules. In both phases of the experiment, adjusting delays in successive blocks of trials were analyzed using the Fourier transform. The power spectrum obtained from individual rats had a dominant frequency that corresponded to a period of oscillation of the adjusting delay between 30 and 100 trial blocks (mean, 78). Power in the dominant frequency band was highest in the early sessions of the first phase and declined with extended training. It is suggested that this experimental protocol may have utility in neurobehavioral studies of intertemporal 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.     

Impulsive choice and pre‐exposure to delays: iv. effects of delay‐ and immediacy‐exposure training relative to maturational changes in impulsivity

Impulsive choice describes preference for smaller, sooner rewards over larger, later rewards. Excessive delay discounting (i.e., rapid devaluation of delayed rewards) underlies some impulsive choices, and is observed in many maladaptive behaviors (e.g., substance abuse, gambling). Interventions designed to reduce delay discounting may provide therapeutic gains. One such intervention provides rats with extended training with delayed reinforcers. When compared to a group given extended training with immediate reinforcers, delay‐exposed rats make significantly fewer impulsive choices. To what extent is this difference due to delay‐exposure training shifting preference toward self‐control or immediacy‐exposure training (the putative control group) shifting preference toward impulsivity? The current study compared the effects of delay‐ and immediacy‐exposure training to a no‐training control group and evaluated within‐subject changes in impulsive choice across 51 male Wistar rats. Delay‐exposed rats made significantly fewer impulsive choices than immediacy‐exposed and control rats. Between‐group differences in impulsive choice were not observed in the latter two groups. While delay‐exposed rats showed large, significant pre‐ to posttraining reductions in impulsive choice, immediacy‐exposed and control rats showed small reductions in impulsive choice. These results suggest that extended training with delayed reinforcers reduces impulsive choice, and that extended training with immediate reinforcers does not increase impulsive choice.     

Preference reversals with food and water reinforcers in rats

Rats were given a choice between a smaller, immediately available reward and a larger reward available after a delay. In one phase, the reward was food and in another phase, the reward was water. Constant delays were added between the choice presentation and the delivery of the reward alternatives. As the time between choice and reward delivery increased from 0 to 25 s, all rats (except one in the water phase) reversed their preference from the smaller, sooner alternative to the larger, later alternative. These findings extend the generality of the preference-reversal animal model to qualitatively different reinforcers. Furthermore, the presence of both impulsive and self-control choices within the same animal is consistent with the view that self-control may be understood as choice behavior, and that species differences in self-control may be differences in degree, not kind.     

RISKY CHOICE IN PIGEONS AND HUMANS: A CROSS‐SPECIES COMPARISON

Pigeon and human subjects were given repeated choices between variable and adjusting delays to token reinforcement that titrated in relation to a subject's recent choice patterns. Indifference curves were generated under two different procedures: immediate exchange, in which a token earned during each trial was exchanged immediately for access to the terminal reinforcer (food for pigeons, video clips for humans), and delayed exchange, in which tokens accumulated and were exchanged after 11 trials. The former was designed as an analogue of procedures typically used with nonhuman subjects, the latter as an analogue to procedures typically used with human participants. Under both procedure types, different variable‐delay schedules were manipulated systematically across conditions in ways that altered the reinforcer immediacy of the risky option. Under immediate‐exchange conditions, both humans and pigeons consistently preferred the variable delay, and indifference points were generally ordered in relation to relative reinforcer immediacies. Such risk sensitivity was greatly reduced under delayed‐exchange conditions. Choice and trial‐initiation response latencies varied directly with indifference points, suggesting that local analyses may provide useful ancillary measures of reinforcer value. On the whole, the results indicate that modifying procedural features brings choices of pigeons and humans into better accord, and that human—nonhuman differences on risky choice procedures reported in the literature may be at least partly a product of procedural differences.     

Stimulus Equivalence: Effects Of A Default-response Option On Emergence Of Untrained Stimulus Relations

Human subjects were exposed to a concurrent-chains schedule in which reinforcer amounts, delays, or both were varied in the terminal links, and consummatory responses were required to receive points that were later exchangeable for money. Two independent variable-interval 30-s schedules were in effect during the initial links, and delay periods were defined by fixed-time schedules. In Experiment 1, subjects were exposed to three different pairs of reinforcer amounts and delays, and sensitivity to reinforcer amount and delay was determined based on the generalized matching law. The relative responding (choice) of most subjects was more sensitive to reinforcer amount than to reinforcer delay. In Experiment 2, subjects chose between immediate smaller reinforcers and delayed larger reinforcers in five conditions with and without timeout periods that followed a shorter delay, in which reinforcer amounts and delays were combined to make different predictions based on local reinforcement density (i.e., points per delay) or overall reinforcement density (i.e., points per total time). In most conditions, subjects' choices were qualitatively in accord with the predictions from the overall reinforcement density calculated by the ratio of reinforcer amount and total time. Therefore, the overall reinforcement density appears to influence the preference of humans in the present self-control choice situation.     

Effects of time between trials on rats' and pigeons' choices with probabilistic delayed reinforcers

Parallel experiments with rats and pigeons examined reasons for previous findings that in choices with probabilistic delayed reinforcers, rats' choices were affected by the time between trials whereas pigeons' choices were not. In both experiments, the animals chose between a standard alternative and an adjusting alternative. A choice of the standard alternative led to a short delay (1 s or 3 s), and then food might or might not be delivered. If food was not delivered, there was an "interlink interval," and then the animal was forced to continue to select the standard alternative until food was delivered. A choice of the adjusting alternative always led to food after a delay that was systematically increased and decreased over trials to estimate an indifference point--a delay at which the two alternatives were chosen about equally often. Under these conditions, the indifference points for both rats and pigeons increased as the interlink interval increased from 0 s to 20 s, indicating decreased preference for the probabilistic reinforcer with longer time between trials. The indifference points from both rats and pigeons were well described by the hyperbolic-decay model. In the last phase of each experiment, the animals were not forced to continue selecting the standard alternative if food was not delivered. Under these conditions, rats' choices were affected by the time between trials whereas pigeons' choices were not, replicating results of previous studies. The differences between the behavior of rats and pigeons appears to be the result of procedural details, not a fundamental difference in how these two species make choices with probabilistic delayed reinforcers.     

Choice in a self-control paradigm: Quantification of experience-based differences

Previous quantitative models of choice in a self-control paradigm (choice between a larger, more-delayed reinforcer and a smaller, less-delayed reinforcer) have not described individual differences. Two experiments are reported that provide additional quantitative data on experience-based differences in choice between reinforcers of varying sizes and delays. In Experiment 1, seven pigeons in a self-control paradigm were exposed to a fading procedure that increased choices of the larger, more-delayed reinforcer through gradually decreasing the delay to the smaller of two equally delayed reinforcers. Three control subjects, exposed to each of the small-reinforcer delays to which the experimental subjects were exposed, but for fewer sessions, demonstrated that lengthy exposure to each of the conditions in the fading procedure may be necessary in order for the increase to occur. In Experiment 2, pigeons with and without fading-procedure exposure chose between reinforcers of varying sizes and delays scheduled according to a concurrent variable-interval variable-interval schedule. In both experiments, pigeons with fading-procedure exposure were more sensitive to variations in reinforcer size than reinforcer delay when compared with pigeons without this exposure. The data were described by the generalized matching law when the relative size of its exponents, representing subjects' relative sensitivity to reinforcer size and delay, were grouped according to subjects' experience.     

DELAY‐AMOUNT TRADEOFFS IN CHOICES BY PIGEONS AND RATS: HYPERBOLIC VERSUS EXPONENTIAL DISCOUNTING

An adjusting‐delay procedure was used to study the choices of pigeons and rats when both delay and amount of reinforcement were varied. In different conditions, the choice alternatives included one versus two reinforcers, one versus three reinforcers, and three versus two reinforcers. The delay to one alternative (the standard alternative) was kept constant in a condition, and the delay to the other (the adjusting alternative) was increased or decreased many times a session so as to estimate an indifference point—a delay at which the two alternatives were chosen about equally often. Indifference functions were constructed by plotting the adjusting delay as a function of the standard delay for each pair of reinforcer amounts. The experiments were designed to test the prediction of a hyperbolic decay equation that the slopes of the indifference functions should increase as the ratio of the two reinforcer amounts increased. Consistent with the hyperbolic equation, the slopes of the indifference functions depended on the ratios of the two reinforcer amounts for both pigeons and rats. These results were not compatible with an exponential decay equation, which predicts slopes of 1 regardless of the reinforcer amounts. Combined with other data, these findings provide further evidence that delay discounting is well described by a hyperbolic equation for both species, but not by an exponential equation. Quantitative differences in the y‐intercepts of the indifference functions from the two species suggested that the rate at which reinforcer strength decreases with increasing delay may be four or five times slower for rats than for pigeons.     

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.     

Females in the forefront: time-based intervention effects on impulsive choice and interval timing in female rats

Impulsive choice has been implicated in substance abuse, gambling, obesity, and other maladaptive behaviors. Deficits in interval timing may increase impulsive choices, and therefore, could serve as an avenue through which suboptimal impulsive choices can be moderated. Temporal interventions have successfully attenuated impulsive choices in male rats, but the efficacy of a temporal intervention has yet to be assessed in female rats. As such, this experiment examined timing and choice behavior in female rats, and evaluated the ability of a temporal intervention to mitigate impulsive choice behavior. The temporal intervention administered in this study was successful in reducing impulsive choices compared to a control group. Results of a temporal bisection task indicated that the temporal intervention increased long responses at the shorter durations. Further, results from the peak trials within the choice task combined with the progressive interval task suggest that the intervention increased sensitivity to delay and enhanced timing confidence. Overall, these results indicate that a temporal intervention can be a successful avenue for reducing impulsive choice behavior in female rats, and could contribute to the development of behavioral interventions to prevent impulsive choice and maladaptive behaviors that can be applied to both sexes.     

Sensitivity to reinforcer duration in a self-control procedure

In a concurrent-chains procedure, pigeons' responses on left and right keys were followed by reinforcers of different durations at different delays following the choice responses. Three pairs of reinforcer delays were arranged in each session, and reinforcer durations were varied over conditions. In Experiment 1 reinforcer delays were unequal, and in Experiment 2 reinforcer delays were equal. In Experiment 1 preference reversal was demonstrated in that an immediate short reinforcer was chosen more frequently than a longer reinforcer delayed 6 s from the choice, whereas the longer reinforcer was chosen more frequently when delays to both reinforcers were lengthened. In both experiments, choice responding was more sensitive to variations in reinforcer duration at overall longer reinforcer delays than at overall shorter reinforcer delays, independently of whether fixed-interval or variable-interval schedules were arranged in the choice phase. We concluded that preference reversal results from a change in sensitivity of choice responding to ratios of reinforcer duration as the delays to both reinforcers are lengthened.     

Effects of arachidonic acid on the spatial cognition of aged rats

Abstract:  To examine the effects of arachidonic acid (AA) on age-related cognitive deficits, Fischer 344 rats were given an AA-supplemented powder diet for 12 weeks from 18.6 months old as part of the old AA (OA) group. Other age-matched rats were fed a non-AA-supplemented powder diet as part of the old control (OC) group, and young rats were fed lab pellets as part of the young control (YC) group. When the aged rats reached 20.0 months old, all animals were tested for spontaneous activity in an open field, followed by the Morris water maze place and cue tasks. Escape latencies for the place task showed that the YC rats were fastest, and there was no difference between the OA and OC groups. However, the probe test and the first block of the cue task indicated that the OA rats remembered the location of an invisible platform better than the OC rats. Although the amount of hippocampal AA was larger in the OA rats than in the OC rats, the difference was not significant. The results showed that AA administration to aged animals may alleviate age-related deficits in spatial cognition.     

Building blocks of self-control: increased tolerance for delay with bundled rewards

Impulsive choice can be defined as temporary preference for a smaller-sooner reward (SS) over a larger-later reward (LL). Hyperbolic discounting implies that impulsive choices will occur less when organisms choose between a series of SSs versus LLs all at once than when they choose between single SS versus LL pairs. Eight rats were exposed to two conditions of an intertemporal choice paradigm using sucrose solution as reward. In both conditions, the LL was 150 microl delayed by 3 s, while the SS was an immediate reward that ranged from 25-150 microl across sessions. Preference for the LL was greater when the chosen reward was automatically delivered three times in succession (bundled) than when it was chosen singly and delivered after each choice. For each of the 8 rats, the estimated SS amount that produced indifference was higher in the bundled condition than in the single condition. Because bundling in humans may be based on the perception that one's current choice is predictive of future choices, the data presented here may demonstrate an important building block of self-control.     

Preference for progressive delays and concurrent physical therapy exercise in an adult with acquired brain injury

The purpose of this study was to increase self-control and engagement in a physical therapy task (head holding) for a man with acquired traumatic brain injury. Once impulsivity was observed (i.e., repeated impulsive choices), an experimental condition was introduced that consisted of choices between a small immediate reinforcer, a large fixed-delay reinforcer, and a large progressive-delay reinforcer. The participant showed a preference for the progressive-delay option, even when the duration of the delay exceeded that of the fixed delay. The results have implications for establishing optimal choice making and teaching life-enhancing skills.