Role of conditioned reinforcers in the initiation, maintenance and extinction of drug-seeking behavior.

The development of a secondary reinforcer as a result of associating a neutral stimulus (buzzer) with intravenous (IV) doses of morpine was studied in rats. Secondary reinforcement developed in the absence of physical dependence and followed the association of the stimulus with either response-contingent or non-contingent injections of morphine. Strength of the conditioned reinforcer, measured in terms of responding on a lever for the stimulus plus infusion of saline solution, was proportional to the unit dosage of morphine employed in pairings of buzzer and drug. When extinction of the lever-press response for IV morphine was conducted (by substituting saline for morphine solution) in the absence of the conditioned reinforcing stimulus, it was seen later that the stimulus could still elicit lever responses, until it too had been present for a sufficient interval of non-reinforced responding. Similarly, extinction of the response for morphine by blocking its action with naloxone in the absence of the stimulus did not eliminate the conditioned reinforcement. Another study showed that a passive, subcutaneous (SC) dose of morphine served to maintain lever-pressing on a contingency of buzzer plus saline infusion. Furthermore, the stimuli resulting from the presence of morphine (after a SC injection) were able to reinstate the lever-responding with only the buzzer-saline contingency when such responses had previously been extinguished. Moreover, it was shown that d-amphetamine could restore responding under the same conditions, and that morphine could also do so for rats in which the primary reinforcer had been d-amphetamine. It is suggested that animal data such as these show that procedures designed for the elimination of human drug-taking behavior must take into account secondary reinforcers as well as the primary reinforcer(s).     

Resistance to extinction and behavioral momentum

In the metaphor of behavioral momentum, reinforcement is assumed to strengthen discriminated operant behavior in the sense of increasing its resistance to disruption, and extinction is viewed as disruption by contingency termination and reinforcer omission. In multiple schedules of intermittent reinforcement, resistance to extinction is an increasing function of reinforcer rate, consistent with a model based on the momentum metaphor. The partial-reinforcement extinction effect, which opposes the effects of reinforcer rate, can be explained by the large disruptive effect of terminating continuous reinforcement despite its strengthening effect during training. Inclusion of a term for the context of reinforcement during training allows the model to account for a wide range of multiple-schedule extinction data and makes contact with other formulations. The relation between resistance to extinction and reinforcer rate on single schedules of intermittent reinforcement is exactly opposite to that for multiple schedules over the same range of reinforcer rates; however, the momentum model can give an account of resistance to extinction in single as well as multiple schedules. An alternative analysis based on the number of reinforcers omitted to an extinction criterion supports the conclusion that response strength is an increasing function of reinforcer rate during training.     

Response-reinforcer relations and resistance to change

Behavioral momentum theory suggests that the relation between a response and a reinforcer (i.e., response-reinforcer relation) governs response rates and the relation between a stimulus and a reinforcer (i.e., stimulus-reinforcer relation) governs resistance to change. The present experiments compared the effects degrading response-reinforcer relations with response-independent or delayed reinforcers on resistance to change in conditions with equal stimulus-reinforcer relations. In Experiment 1, pigeons responded on equal variable-interval schedules of immediate reinforcement in three components of a multiple schedule. Additional response-independent reinforcers were available in one component and additional delayed reinforcers were available in another component. The results showed that resistance to disruption was greater in the components with added reinforcers than without them (i.e., better stimulus-reinforcer relations), but did not differ for the components with added response-independent and delayed reinforcement. In Experiment 2, a component presenting immediate reinforcement alternated with either a component that arranged equal rates of reinforcement with a proportion of those reinforcers being response independent or a component with a proportion of the reinforcers being delayed. Results showed that resistance to disruption tended to be either similar across components or slightly lower when response-reinforcer relations were degraded with either response-independent or delayed reinforcers. These findings suggest that degrading response-reinforcer relations can impact resistance to change, but that impact does not depend on the specific method and is small relative to the effects of the stimulus-reinforcer relation.     

Four-alternative choice violates the constant-ratio rule

Six pigeons responded on a four-key concurrent variable-interval schedule in which a 27:9:3:1 distribution of reinforcers between the keys changed every 10 reinforcers. Their behaviour quickly came under the control of this changing four-way reinforcer ratio. However, preference between a pair of keys depended not only on the relative reinforcer rates on those keys, but also on the absolute levels of those rates. This contradicts the constant-ratio rule that underpins the matching approach to choice, but is predicted by a contingency-discriminability model that assumes that organisms may occasionally misattribute reinforcers to a response that did not produce them. Reinforcers produced strong preference pulses, or transient increases in responding on the just-reinforced key. Despite accurate tracking of the reinforcer ratio, reinforcers obtained late in components and from leaner keys still produced strong pulses, suggesting both extended and local control of behaviour. Patterns of switching between keys were graded and similarly controlled by the reinforcer rates on each key. Whether considered in terms of switching, local preference pulses, or extended preference, behaviour was controlled by a rapidly changing four-way reinforcer ratio in a graduated, continuous manner that is unlikely to be explained by a simple heuristic such as fix-and-sample.     

Haloperidol, dynamics of choice, and the parameters of the matching law

The idea that dopamine mediates the reinforcing effects of stimuli persists in the field of neurosciences. The present study shows that haloperidol, a dopamine antagonist, does not eliminate the reinforcing value of food reinforcers. The ratio of reinforcers changed seven times across two levers within sessions, modeling a dynamic environment. The magnitude of the reinforcer was manipulated independently of the reinforcer ratio. Four doses of intraperitoneal haloperidol were assessed over periods of 12 daily sessions. Haloperidol did not impair the discrimination that the rats established between rich and lean levers; the response distributions favored the lever associated with the higher probability of reinforcement and the larger pellets. The parameters of the generalized matching law (bias and sensitivity) were used to estimate effects of haloperidol upon the motor system and upon the rats' motivation for food reinforcers.     

Delay discounting of saccharin in rhesus monkeys

The value of a reinforcer decreases as the time until its receipt increases, a phenomenon referred to as delay discounting. Although delay discounting of non-drug reinforcers has been studied extensively in a number of species, our knowledge of discounting in non-human primates is limited. In the present study, rhesus monkeys were allowed to choose in discrete trials between 0.05% saccharin delivered in different amounts and with different delays. Indifference points were calculated and discounting functions were established. Discounting functions for saccharin were well described by a hyperbolic function. Moreover, the discounting rates for saccharin in all six monkeys were comparable to those of other non-human animals responding for non-drug reinforcers. Also consistent with other studies of non-human animals, changing the amount of a saccharin reinforcer available after a 10-s delay did not affect its relative subjective value. Discounting functions for saccharin were steeper than we found in a previous study with cocaine, raising the possibility that drugs such as cocaine may be discounted less steeply than non-drug reinforcers.     

Human performance on a two-alternative rapid-acquisition choice task

Davison and Baum [Davison, M., Baum, W. M., 2000. Choice in a variable environment: every reinforcer counts. Journal of the Experimental Analysis of Behavior 74, 1-24.] developed a concurrent-schedule procedure where, within each session, different reinforcer ratios were arranged across components separated by brief black-outs. Behaviour adapted quickly to the reinforcer ratios and reinforcers also had local effects on responding. This procedure has been used with pigeons and rats. In the present experiment, we adapted the Davison and Baum procedure to study the effects of reinforcement on human choice behaviour. Eighteen participants were presented with four different reinforcer ratios within a single 50-minute session. Mean sensitivity to the reinforcer ratios increased within components, and preference was greater for the just-reinforced response alternative immediately following reinforcer delivery, similar to the results from non-human experiments. Although there were limitations to the current procedure, the local time scale analyses are a novel way of examining human operant behaviour.     

Mathematical principles of reinforcement and resistance to change

Although Killeen's mathematical principles of reinforcement (MPR) apply to the asymptotic rate of a free operant after extended exposure to a single schedule of reinforcement, they can be extended to resistance to change in multiple schedules via alterations in the parameter representing the activating effects of reinforcers. MPR's predictions of resistance to change in relation to reinforcer rate on variable-interval (VI) schedules are empirically correct and agree with behavioral momentum theory (BMT). However, both MPR and BMT encounter problems in accounting for the effects of delayed reinforcement on resistance to change, relative to immediate reinforcement at the same rate. Further problems are raised by differences in resistance to change between variable-ratio (VR) and variable-interval performances maintained by the same reinforcer rate. With both delayed versus immediate reinforcement and variable-ratio versus variable-interval reinforcement, differential resistance to change is negatively correlated with the log ratios of baseline response rates when reinforcer rates are equated. Cases where resistance to change varies despite equated reinforcer rates, and the correlations among behavioral measures, provide challenges and opportunities for both MPR and BMT.     

Reinforcer satiation and resistance to change of responding maintained by qualitatively different reinforcers

In previous research on resistance to change, differential disruption of operant behavior by satiation has been used to assess the relative strength of responding maintained by different rates or magnitudes of the same reinforcer in different stimulus contexts. The present experiment examined resistance to disruption by satiation of one reinforcer type when qualitatively different reinforcers were arranged in different contexts. Rats earned either food pellets or a 15% sucrose solution on variable-interval 60-s schedules of reinforcement in the two components of a multiple schedule. Resistance to satiation was assessed by providing free access either to food pellets or the sucrose solution prior to or during sessions. Responding systematically decreased more relative to baseline in the component associated with the satiated reinforcer. These findings suggest that when qualitatively different reinforcers maintain responding, relative resistance to change depends upon the relations between reinforcers and disrupter types.     

Contingency discriminability and the generalized matching law describe choice on concurrent ratio schedules of wheel-running reinforcement

Belke (2010) showed that on concurrent ratio schedules, the difference in ratio requirements required to produce near exclusive preference for the lower ratio alternative was substantively greater when the reinforcer was wheel running than when it was sucrose. The current study replicated this finding and showed that this choice behavior can be described by the matching law and the contingency discriminability model. Eight female Long Evans rats were exposed to concurrent VR schedules of wheel-running reinforcement (30s) and the schedule value of the initially preferred alternative was systematically increased. Two rats rapidly developed exclusive preference for the lower ratio alternative, but the majority did not - even when ratios differed by 20:1. Analysis showed that estimates of slopes from the matching law and the proportion of reinforcers misattributed from the contingency discriminability model were related to the ratios at which near exclusive preference developed. The fit of these models would be consistent with misattribution of reinforcers or poor discrimination between alternatives due to the long duration of wheel running.     

Varying reinforcer duration produces behavioral interactions during multiple schedules

The experiments tested the idea that changes in habituation to the reinforcer contribute to behavioral interactions during multiple schedules. This idea predicts that changing an aspect of the reinforcer should disrupt habituation and produce an interaction. Pigeons and rats responded on multiple variable interval variable interval schedules. Introducing variability into the duration of reinforcers in one component increased response rates in both components when the schedules provided high, but not low, rates of reinforcement. The increases in constant-component response rates grew larger as the session progressed. Within-session decreases in responding were smaller when the other component provided variable-, rather than fixed-, duration reinforcers. These results are consistent with the idea that changes in habituation to the reinforcer contribute to behavioral interactions. They help to explain why interactions do not occur for some subjects under conditions that produce them for others. Finally, the results question the assumption that induction and behavioral contrast are always produced by different theoretical mechanisms.     

The operant reserve: a computer simulation in (accelerated) real time

In Skinner's Reflex Reserve theory, reinforced responses added to a reserve depleted by responding. It could not handle the finding that partial reinforcement generated more responding than continuous reinforcement, but it would have worked if its growth had depended not just on the last response but also on earlier responses preceding a reinforcer, each weighted by delay. In that case, partial reinforcement generates steady states in which reserve decrements produced by responding balance increments produced when reinforcers follow responding. A computer simulation arranged schedules for responses produced with probabilities proportional to reserve size. Each response subtracted a fixed amount from the reserve and added an amount weighted by the reciprocal of the time to the next reinforcer. Simulated cumulative records and quantitative data for extinction, random-ratio, random-interval, and other schedules were consistent with those of real performances, including some effects of history. The model also simulated rapid performance transitions with changed contingencies that did not depend on molar variables or on differential reinforcement of inter-response times. The simulation can be extended to inhomogeneous contingencies by way of continua of reserves arrayed along response and time dimensions, and to concurrent performances and stimulus control by way of different reserves created for different response classes.     

Human choice behaviour in a frequently changing environment

The present experiment provided a replication in humans of an experimental procedure that has been used frequently with nonhumans to investigate choice behaviour in a changing environment. Six volunteers played a computer game, which required tracking of a moving balloon on two simultaneously available response panels for monetary reinforcers. Each of the 15 sessions randomly arranged the following concurrent variable-interval reinforcement schedules, which were in effect until six reinforcers had been obtained: 27:1, 9:1, 3:1, 1:1, 1:3, 1:9, and 1:27. Although many aspects of human performance appeared to be qualitatively similar to that of nonhumans on this procedure, such as the rapid preference shifts towards the within-session reinforcer ratios and the presence of local effects of reinforcers, values of sensitivity to reinforcement were very variable in the present study, as commonly reported in human choice studies. Future variations and refinements of the experimental methods are needed to explore how this variability may be reduced.     

Small and large fixed ratios with the same unit price

Key pressing of rats was maintained under multiple and discrete-trial choice schedules with reinforcer units of 45 mg food pellets or 3.5 s dips of sucrose solution. Both smaller and larger fixed ratio (FR) schedules were associated with the same unit price in a manner, for example, that each of eight iterations of FR120 was associated with delivery of a single reinforcer unit and one instance of FR960 was associated with eight reinforcer units. FR requirement varied between 20 and 1560 per aggregate reinforcer and unit price varied between 20 and 240 per reinforcer unit. During multiple schedules with food reinforcers, rates and patterns of responding were comparable over nearly a 50-fold range of FR requirements (20-1380) when unit price was 20; over nearly a six-fold range of FR requirements (120-720) when unit price was 120; and was only marginally maintained when unit price was 240. Demand for food pellets was comparatively inelastic at FRs between 20 and 120, during which subjects did not receive supplemental feeding outside experimental sessions, but was elastic at FRs greater than 240, when subjects sometimes did receive supplemental feeding. In a discrete-trial choice procedure with a constant unit price of 120 for sucrose solution, subjects were indifferent between smaller FRs and alternative FRs as large as 480, but began switching away from larger FRs that were 600 or greater. Because responding had been comparably maintained under both FR120 and FRs as large as 960 in the multiple schedule, results from the choice procedure indicated that choice performance was influenced by variables other than FR requirement and unit price. Because aggregate reinforcers were the same for smaller and larger FRs, the most likely reason for preferring smaller FRs was the nearness in time to some reinforcer.     

Species differences between rats and pigeons in choices with probabilistic and delayed reinforcers

An adjusting-delay procedure was used to study rats' choices with probabilistic and delayed reinforcers, and to compare them with previous results from pigeons. A left lever press led to a 5-s delay signaled by a light and a tone, followed by a food pellet on 50% of the trials. A right lever press led to an adjusting delay signaled by a light followed by a food pellet on 100% of the trials. In some conditions, the light and tone for the probabilistic reinforcer were present only on trials that delivered food. In other conditions, the light and tone were present on all trials that the left lever was chosen. Similar studies with pigeons [Mazur, J.E., 1989. Theories of probabilistic reinforcement. J. Exp. Anal. Behav. 51, 87-99; Mazur, J.E., 1991. Conditioned reinforcement and choice with delayed and uncertain primary reinforcers. J. Exp. Anal. Behav. 63, 139-150] found that choice of the probabilistic reinforcer increased dramatically when the delay-interval stimuli were omitted on no-food trials, but this study found no such effect with the rats. In other conditions, the probability of food was varied, and comparisons to previous studies with pigeons indicated that rats showed greater sensitivity to decreasing reinforcer probabilities. The results support the hypothesis that rats' choices in these situations depend on the total time between a choice response and a reinforcer, whereas pigeons' choices are strongly influenced by the presence of delay-interval stimuli.     

Behavioral economics of drug self-administration. I. Functional equivalence of response requirement and drug dose

Response requirement and dose of drug per administration are two separate factors that have been demonstrated to control drug self-administration. Recent developments in behavioral economics have shown that these two factors are in fact functionally equivalent for nondrug reinforcers, as indicated by a unit-price analysis. In this review, the unit-price notion was tested for drugs as reinforcers via a re-analysis of ten drug self-administration studies. The results of the re-analysis indicated that response requirement and reinforcer magnitude, the constituents of unit price, have functionally equivalent effects on drug consumption and that a positively decelerating demand curve is produced as unit price increases. This suggests that the behavioral-economic notion of unit price is a more parsimonious explanation of the effects of response requirement and dose in drug self-administration studies, in that it integrates and describes what was previously considered to be two distinct operations.     

The role of place of reinforcer delivery in the appearance of positive induction when rats respond for 1% sucrose

Rats increase their rate of operant responding for 1% sucrose reinforcement in the first half of an experimental session if a high-valued reinforcer will be available in the second half. Previous research suggests that this induction effect occurs because the reinforcing value of the low-valued substance has increased. The present study investigated whether this increase may occur because of where the substances are delivered. Rats pressed a lever to earn 1% liquid-sucrose reinforcers in the first half of the session. In control conditions, they also pressed for 1% sucrose in the second half. In treatment conditions, they pressed for food-pellet (Experiment 1) or 32% sucrose (Experiment 2) reinforcers in the second half, with these reinforcers either being delivered to the same location as the 1% sucrose or to a different location. Upcoming food-pellet or 32% sucrose reinforcement increased rates of lever pressing for 1% sucrose in the first half of the session, with the largest increase observed when the high-valued reinforcer was delivered to the same location as the 1% sucrose. Qualitatively similar results were found with rates of consumption of 1% sucrose reinforcers in the first half of the session, which were measured in Experiment 2. The location to which reinforcers are delivered appears to be one of the factors that contributes to this induction effect. The present results may therefore identify one of the factors that determine whether differential conditions of reinforcement will lead to contrast or induction.     

Amount-dependent temporal discounting?

Amount-dependent temporal discounting has been demonstrated for human choice between outcomes differing in amount and delay. In the only study to date with non-humans, Grace reported no evidence for amount-dependent temporal discounting with pigeons in a concurrent-chains procedure. The present experiments repeated Grace's procedure but with modifications to enhance the discrimination between small and large magnitude outcomes. In Experiment 1, sensitivity of pigeons' initial-link choice to the terminal link delay ratio was greater with large reinforcer durations in the terminal links than with small reinforcer durations. This result is consistent with a greater rate of temporal discounting for larger reinforcers (the reverse of the result for humans), but can also be explained as enhanced discrimination of delay ratios with larger reinforcer durations. The results of a second experiment supported Grace's conclusion that amount-dependent temporal discounting does not characterize pigeons' choice in concurrent chains. Because reinforcer amount was held constant between choice alternatives in the present experiments and that of Grace, but varied in the human studies, our results question whether prior demonstrations of amount-dependent discounting reflect the effects of reinforcer delay or of reinforcer amount. Differences in the procedures used to study discounting in humans (titration procedures) and non-humans (concurrent chains) may contribute to the divergent results across species.     

Investigations of timing during the schedule and reinforcement intervals with wheel-running reinforcement

Across two experiments, a peak procedure was used to assess the timing of the onset and offset of an opportunity to run as a reinforcer. The first experiment investigated the effect of reinforcer duration on temporal discrimination of the onset of the reinforcement interval. Three male Wistar rats were exposed to fixed-interval (FI) 30-s schedules of wheel-running reinforcement and the duration of the opportunity to run was varied across values of 15, 30, and 60s. Each session consisted of 50 reinforcers and 10 probe trials. Results showed that as reinforcer duration increased, the percentage of postreinforcement pauses longer than the 30-s schedule interval increased. On probe trials, peak response rates occurred near the time of reinforcer delivery and peak times varied with reinforcer duration. In a second experiment, seven female Long-Evans rats were exposed to FI 30-s schedules leading to 30-s opportunities to run. Timing of the onset and offset of the reinforcement period was assessed by probe trials during the schedule interval and during the reinforcement interval in separate conditions. The results provided evidence of timing of the onset, but not the offset of the wheel-running reinforcement period. Further research is required to assess if timing occurs during a wheel-running reinforcement period.     

The effects of changes in consequences on hens' performance in delayed-matching-to-sample tasks

A delayed-matching-to-sample (DMTS) task was used to investigate remembering with domestic hens. In Conditions 1 and 3 of Experiment 1, six hens responded under a mixed-delay procedure with delays of 0.25, 2, and 8 s. In Condition 2, the reinforcer for correct responding was delayed for 6 s after each correct matching response on 2-s delay trials. In Condition 1, discrimination performance decreased monotonically over the three delays. With the delay to the reinforcer, the decreases were non-monotonic as a result of the considerable drop in the accuracy of discrimination on the 2-s delay trials. Performance at the 2-s delay did not recover completely in Condition 3. In Conditions 1 and 3 of Experiment 2, five hens responded under a mixed-delay procedure with delays of 0, 4, and 16 s. In Condition 2 no reinforcers were provided for correct responding on 0- and 16-s delay trials. When reinforcers were available on all trials discrimination performance decreased monotonically with delay. There were non-monotonic changes in discrimination with delay when there was extinction at two delays resulting mainly from a large drop in discrimination performance at 0 s. In addition, response latencies increased markedly at the two delays associated with extinction. Performance recovered completely in Condition 3. The data support the ideas that remembering involves a temporal discrimination that the effects of delaying reinforcement and removing reinforcement may differ, and that the measurement of response latencies may be a useful tool in DMTS procedures.