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The relationship between reinforcement and memory: parallels in the rewarding and mnemonic effects of the neuropeptide substance P.

A theory of reinforcement is presented which accounts for the backward action of a reinforcer on operant behavior in terms of its effect on memory traces left by the operant. Several possible ways in which a reinforcer could strengthen the probability of recurrence of an operant are discussed. Predictions from the model regarding general memory-promoting effects of reinforcers presented posttrial in various learning paradigms are outlined. The theory also predicts a parallelism in reinforcing and memory-promoting effects of stimuli, including drugs. The second part of the chapter outlines experiments investigating memory modulating and reinforcing effects of the neuropeptide substance P. In general, injection of SP is positively reinforcing when injected into parts of the brain where it has been shown to facilitate learning. Peripheral injection of SP is also reinforcing at the dose known to promote passive avoidance learning when presented posttrial.

Amino Acid Sequence↗

Reinforcing effects of D2 dopamine receptor agonists and partial agonists in rhesus monkeys.

Dopamine (DA) receptors play a role in the reinforcing effects of psychomotor stimulants and other drugs. Both D1 and D2 DA receptor agonists have been reported to function as positive reinforcers in maintaining self-administration in non-human subjects. The purpose of the present study was to evaluate, in monkeys, the reinforcing effects of DA D2 receptor agonists that vary in their efficacy as D2 agonists. Rhesus monkeys were prepared with venous catheters and lever pressing was maintained by i.v. cocaine (n=5, 0.03 mg/kg/inj) in daily baseline sessions (2 h/day, fixed ratio 25). Various doses of cocaine or D2 agonists were then made available for at least four to seven sessions, and until responding was stable. At least one dose of the higher-efficacy D2 agonists, R(-)-propylnorapomorphine (NPA) (n=4, 0.001-0.01 mg/kg/inj), R(-)-apomorphine (APO) (n=4, 0.003-0.1 mg/kg/inj) and R(+)-3-(3-hydroxyphenyl)-N-propylpiperidine HCl [R(+)-3-PPP] (n=4, 0.03-0.3 mg/kg/inj), functioned as a positive reinforcer in all the monkeys tested. In contrast, no dose of the lower-efficacy D2 agonists, R(+)-terguride (n=4, 0.001-0.3 mg/kg/inj) and S(-)-3-(3-hydroxyphenyl)-N-propylpiperidine HCl [S(-)-3-PPP] (n=4, 0.001-0.3 mg/kg/inj), maintained self-administration. In in vitro binding studies with monkey brain tissue NPA and terguride had high affinities for the D2 receptor, while APO had intermediate affinity, and the 3-PPPs had low affinity. Among the compounds that were reinforcers potency as a reinforcer was directly related to D2 affinity in three of the four monkeys, consistent with D2 receptor involvement in the reinforcing effect of these compounds. The data suggest that the efficacy at D2 receptors is directly related to the reinforcing effect.

Animals↗

Relative reinforcing effects of different benzodiazepine doses for rhesus monkeys.

The relative reinforcing effects of different doses of benzodiazepines were determined by giving rhesus monkeys concurrent access to different diazepam and midazolam concentrations. For each monkey a dose response function was obtained using three drug concentrations: low (L), intermediate (I), and high (H). The benzodiazepine and the water vehicle were concurrently available under independent fixed-ratio (FR) schedules. After establishing that each concentration was a reinforcer in comparison to vehicle, relative preference for the different concentrations was examined by making pairs of concentrations concurrently available under independent FR schedules. Three pairs were studied (H vs. L, H vs. I, and I vs. L). With both drugs, higher concentrations maintained greater response rates than lower concentrations. Thus, relative reinforcing effects increased with dose. These findings are similar to those obtained with other reinforcing drugs and provide further evidence that benzodiazepines share significant characteristics with other drug reinforcers. Importantly, absolute response rates (responses per session) obtained when only one drug dose was present were not reliably predictive of subsequent preferences for the dose. Both benzodiazepines served as effective reinforcers in that consistent levels of responding were maintained across doses and above vehicle levels under concurrent FR 32 schedules. As with other reinforcing drugs, the reinforcing effects of benzodiazepines increase with increases in dose over a broad range of values.

Administration, Oral↗

A within-subject comparison of three different schedules of reinforcement of drug abstinence using cigarette smoking as an exemplar.

In this experiment we compared three different schedules of reinforcement for promoting and sustaining short-term drug abstinence. For pragmatic reasons, cigarette smoking was studied as an exemplar of drug self-administration. The three schedules studied were a fixed magnitude of reinforcement for abstinence, a progressive increase in magnitude of reinforcement for abstinence with a reset contingency for drug use, and a progressive increase in magnitude of reinforcement for abstinence without a reset contingency. Eighteen cigarette smokers experienced the three schedules in a counterbalanced order. Each schedule was in effect for 5 consecutive days (M-F), during which time abstinence was reinforced according to the different schedules of reinforcement. The total amount of reinforcement (money) available was the same during each condition. The progressive magnitude with a reset schedule was more effective than the other two schedules in sustaining an initial period of abstinence. These results systematically replicate and extend those from prior studies demonstrating the efficacy of schedules incorporating a progressively increasing magnitude of reinforcement with a reset contingency for sustaining initial drug abstinence, and demonstrate the importance of the reset contingency to that effect.

Adult↗

The impact of reinforcement density on response differentiation in configural discrimination problems.

Two human Pavlovian conditioning experiments investigated the impact of reinforcement density (the number of reinforced trials divided by the total number of trials) on discrimination learning. Experiment 1 used a negative patterning problem (A+, B+, AB-) and Experiment 2 used a positive patterning problem (A-, B-, AB+). In both experiments, reinforcement density varied across four levels. Response differentiation between reinforced and non-reinforced stimuli was linearly related to the decrease in reinforcement density. Responses to nonreinforced stimuli did not differ between the four groups in either experiment. In contrast to this, responses to reinforced stimuli were considerably more pronounced in conditions with lower reinforcement density. For negative patterning, this replicates and extends similar observations from other species. For positive patterning, this is a finding that has not yet been reported in other experiments. The results are in agreement with the comparator hypothesis (Miller & Matzel, 1988) and with Wagner's (1981) "standard operating procedures" (SOP) model.

Adult↗

The representation of the reinforcer and the force of the pigeon's keypeck in first- and second-order conditioning.

The aim of this study was to determine whether reinforcer-specific conditioned responding would occur in a situation in which responding was not thought to be mediated by a representation that encodes information about the specific properties of the reinforcer. The force of the pigeon's keypeck was monitored during first- and second-order conditioning with either food or water as the unconditioned stimulus (US). Each pigeon was trained with four different stimuli: a first-order cue predicting that responding would be reinforced with grain (S1f), a first-order cue predicting water as the reinforcer (S1w), a second-order stimulus predicting the S1f (S2f), and a second-order cue predicting the S1w (S2w). Following conditioning, the pigeons were selectively satiated with one of the two reinforcers and presented with the first- and second-order cues in an extinction test. At the end of training, the pigeon's keypecks were less forceful to the S1w than to the S1f. There was not, however, a reliable difference between the force of the pecks to the S2f and the S2w. These force differences are consistent with the conclusion that the topography of the keypecks was systematically related to the nature of the primary reinforcer during first- but not during second-order conditioning. The results from the selective satiation test are difficult to interpret. There was no evidence to indicate that second-order responding was mediated by a detailed representation of the primary US, but a detailed representation of the reinforcer may have been mediating first-order responding. Taken together, these findings are consistent with the view that a representation of the reinforcer is an important determinant of the topography of conditioned responding.

Animals↗

The effect of reinforcement variables on inhibition in children with ADHD.

Behavioral inhibition, often cited as a central deficit in children with ADHD, has been shown to change in response to reinforcement. In this preliminary investigation, children with ADHD (n= 20) and matched controls (ages 7 to 12) completed a new version of the stop signal paradigm, the Fire Fighter Game, a measure of inhibition of a prepotent motor response, under three conditions: (1) no reinforcement; (2) immediate reinforcement; and (3) delayed reinforcement. In all conditions, the stop signal reaction time (SSRT) of children with ADHD was consistently longer than controls. Both groups improved significantly with reinforcement, and there was no strong evidence that immediate reinforcement was more effective than delayed reinforcement. However, it appeared that the reason for the changes in SSRT in response to reinforcement differed between the groups. Children in the control group responded faster on go trials, whereas children with ADHD improved their ability to inhibit responding with shorter stop delays. The relevance of these findings is discussed in terms of current theories of ADHD.

Analysis of Variance↗

Prenatal ethanol exposure increases ethanol reinforcement in neonatal rats.

BACKGROUND: Recent experimental findings indicate that newborn rats are quite susceptible to the reinforcing properties of ethanol (EtOH). The basis seems to be more pharmacological than gustatory, olfactory, or caloric as a single pairing of a surrogate nipple providing water with an intraperitoneal (i.p.) injection of EtOH results in conditioned attachment to an empty surrogate nipple. The present study tested whether prenatal exposure to moderate EtOH during late gestation would alter the neonatal rat's susceptibility to this source of EtOH reinforcement and/or EtOH metabolism. METHODS: Pregnant females were given 1 g/kg EtOH or water intragastrically on gestational days (GD) 17 to 20 or were untreated. Rat pups delivered by cesarean section on GD 21 were exposed 3 to 4 hours later to a surrogate nipple providing water, either immediately following or 10 minutes before an i.p. injection of 0.0 (saline), 0.25, 0.5, or 0.75 g/kg EtOH (12.6%, v/v). Ethanol reinforcement was indicated by a significant increase in time attached to an empty surrogate nipple compared with unpaired and saline controls. Testing was conducted 90 minutes after conditioning took place. Blood ethanol levels (BELs) were also taken from subjects injected with identical doses of i.p. EtOH at 5 or 95 minutes after injection. RESULTS: Initial response to a surrogate nipple with water did not differ between groups. For neonates prenatally exposed to EtOH, significant reinforcement by all 3 doses of i.p. EtOH (0.25, 0.5, and 0.75) was found, whereas only the dose of 0.5 g/kg EtOH was reinforcing for neonates in the prenatal water condition and only 0.25 g/kg was reinforcing for pups from untreated dams. Blood ethanol levels after conditioning also differed as a function of prenatal treatment. CONCLUSIONS: These results suggest that prenatal EtOH exposure increases the range of EtOH doses capable of reinforcing effects. Intubations with water, however, eliminated the reinforcing strength of the lowest dose (0.25 g/kg) and shifted the dose-response curve to the right. This effect may be due to the stress of the intubation procedure. Ethanol metabolism was also affected by prenatal EtOH exposure. Pups derived from EtOH-treated dams had slower elimination of EtOH from the blood than all other groups at both the 0.5 and 0.75 g/kg i.p. doses.

Alcohol Drinking↗

Phenobarbitone and the relationship between extraversion and reinforcement in verbal operant conditioning.

The present investigation was designed to study the relationship between phenobarbitone, extraversion and modes of reinforcement in verbal operant conditioning. A factorial design involving three treatments (placebo and two doses, 30 and 100 mg, of phenobarbitone), two levels of extraversion (extraverts and introverts) and two reinforcement conditions ('good' and 'electric shock') with repeated measures on the first factor, was used. Sixty postgraduate female students were individually subjected to Taffel's verbal conditioning procedure. The results support the following conclusions: (1) under the placebo condition, the extraverted subjects condition better with the rewarding reinforcer ('good') while the introverted subjects condition better with the punishing reinforcer ('electric shock'); (2) under the influence of the smaller dose of the drug, as compared with the placebo, the conditioning scores of extraverted subjects increase with the punishing reinforcer but decrease with the rewarding reinforcer, while the conditioning scores of the introverted subjects decrease with the punishing reinforcer, under the influence of the larger dose of the drug, but remain uninfluenced with the rewarding reinforcer.

Adult↗

Response-reinforcer relationships and improved learning in autistic children.

In order to affect more rapid response acquisition for autistic children, researchers have recently begun to investigate the functional relationships of reinforcers to other components of the operant conditioning paradigm. Previous research suggested that functional relationships between target behaviors and reinforcers might be especially effective. For example, locating a reward inside a container might be a more efficient way to teach a child to open the container than by handling the child a reward for opening an empty container. The present experiment assessed, within a multiple baseline design, the possibility of improving autistic children's learning by changing arbitrary response-reinforcer relationships (while holding target behaviors and reinforcers constant) so that the target behaviors became functional (i.e., a direct part of the response chain required for the child to procure the reinforcer). The results showed that: (1) arranging functional response-reinforcer relationships produced immediate improvement in the children's learning, and resulted in rapid acquisition of criterion level responding; and (2) high levels of correct responding initially produced by functional response-reinforcer relationships were continued even when previously ineffective arbitrary response-reinforcer conditions were reinstated. The results are discussed in terms of understanding and improving autistic children's learning.

Autistic Disorder↗

On the relative reinforcing effects of choice and differential consequences.

Research on the reinforcing effects of providing choice-making opportunities to individuals with developmental disabilities (i.e., allowing them to choose reinforcers or tasks) has produced inconsistent results, perhaps because the mechanisms underlying such effects remain unclear. Choice may produce a reinforcement effect because it is correlated with differential consequences (i.e., choice may increase one's access to higher preference stimuli), or it may have reinforcement value independent of (or in addition to) the chosen stimulus. In Experiment 1, we used a concurrent-operants arrangement to assess preference for a choice condition (in which participants selected one of two available reinforcers) relative to a no-choice condition (in which the therapist selected the same reinforcers on a yoked schedule). All 3 participants preferred the choice option. In Experiment 2, we altered the schedules so that the participant selected one of two lower preference reinforcers in the choice condition, whereas the therapist selected a higher preference stimulus for the participant either half or all of the time in the no-choice condition. Participants typically allowed the therapist to select reinforcers for them (i.e., they allocated responding to the no-choice condition) when it resulted in greater access to higher preference stimuli.

Adolescent↗

The measurement and reinforcement of behavior of psychotics.

An attempt was made to strengthen behaviors of psychotics by applying operant reinforcement principles in a mental hospital ward. The behaviors studied were necessary and/or useful for the patient to function in the hospital environment. Reinforcement consisted of the opportunity to engage in activities that had a high level of occurrence when freely allowed. Tokens were used as conditioned reinforcers to bridge the delay between behavior and reinforcement. Emphasis was placed on objective definition and quantification of the responses and reinforcers and upon programming and recording procedures. Standardizing the objective criteria permitted ward attendants to administer the program. The procedures were found to be effective in maintaining the desired adaptive behaviors for as long as the procedures were in effect. In a series of six experiments, reinforced behaviors were considerably reduced when the reinforcement procedure was discontinued; the adaptive behaviors increased immediately when the reinforcement procedure was re-introduced.

Adult↗

Independence of stimulus discriminability from absolute rate of reinforcement in a signal-detection procedure.

Three experiments are reported in which two pigeons were trained to detect differences in stimulus duration under varying levels of absolute rate of reinforcement. Two red stimuli, differing in duration, were arranged probabilistically on the center key of a three-key chamber. On completion of the center-key duration, the center keylight was extinguished and the two side keys were illuminated white. Correct responses were left-key pecks following the shorter duration and right-key pecks following the longer duration. In Experiment 1, relative rate of reinforcement for correct responses was held constant and absolute rate of reinforcement was varied in seven conditions from continuous reinforcement to a variable-interval 90-second schedule. In Experiment 2, relative rate of reinforcement was manipulated across three different absolute rates of reinforcement (continuous reinforcement, variable-interval 15-second, and variable-interval 45-second). Stimulus discriminability was unaffected by changes in absolute or relative rates of reinforcement. Experiment 3 showed that discriminability was also unaffected by arranging the same consequences (three-second blackout) for unreinforced correct responses and errors.

Animals↗

Matching, maximizing, and the behavioral unit: concurrent reinforcement of response sequences.

Pigeons pecked two keys in a probability matching situation in which four two-peck sequences were intermittently reinforced: left-left, left-right, right-left and right-right. In Phase 1, relative reinforcement rate was varied with respect to the first response of a sequence: reinforcers were differentially assigned for left-left and left-right sequences as opposed to right-left and right-right sequences. The second response of reinforced sequences occurred equally on the left and right keys across conditions. In Phase II, relative reinforcement rate was varied for sequences that involve an alternation as opposed to those that did not. The relative outputs of the different sequences matched the relative reinforcement rates for the different sequences in both phases. Relative response rates for key pecks did not always match relative reinforcement rates. The intertrial interval separating responses was varied in both phases; increases in the intertrial interval affected the relative frequency of different sequences. The results demonstrate that response sequences acted as functional units influencing choice and thus support a structural account of choice. At the same time, the matching of relative sequence proportion and relative reinforcement rate supports a matching account.

Animals↗

Control over response number by a targeted percentile schedule: reinforcement loss and the acute effects of d-amphetamine.

Two fixed-consecutive-number-like procedures were used to examine effects of acute d-amphetamine administration on control over response number. In both procedures, rats were required to press the left lever at least once and then press the right lever to complete a trial. The consecutive left-lever presses on each trial comprised a "run." Under the targeted percentile schedule, reinforcement was provided if the current run length was closer to the target length (16) than half of the most recent 24 runs. This differentially reinforced run length while holding reinforcement probability constant at .5. A second group acquired the differentiation under the targeted percentile schedule, but were then shifted to a procedure that yoked reinforcement probability by subject and run length to that obtained under the targeted percentile schedule. The two procedures generated practically identical control run lengths, response rates, reinforcement probabilities, and reinforcement rates. Administration of d-amphetamine disrupted percentile responding to a greater degree than yoked control responding. This disruption decreased reinforcement frequency less in the former than the latter procedure. The similar baseline responding under these two procedures suggests that this difference in sensitivity was due to behavioral adjustments to drug prompted by reduction of reinforcement density in the yoked control but not the percentile schedule. These adjustments attenuate the drug's effects under the former, but not the latter, procedure.

Animals↗

Acquisition of a spatially defined operant with delayed reinforcement.

Two experiments investigated the role of an immediate, response-produced auditory stimulus during acquisition, via delayed reinforcement, of a response selected to control for possible unprogrammed, operandum-related sources of response feedback. Experimentally naive rats were exposed to a delayed-food reinforcement condition, specifically a tandem fixed-ratio 1 differential-reinforcement-of-other-behavior 30-s schedule. The response was defined as breaking a photocell beam located near the ceiling at the rear of the operant conditioning chamber. In Experiment 1, rates of photobeam breaking by each rat increased from near zero, regardless of the presence or absence of a tone that immediately followed the response initiating the delay interval. Though not essential, the tone facilitated response acquisition and resulted in more efficient response patterns at stability. Experiment 2 demonstrated that photobeam-breaking response rates under the delayed reinforcement contingency exceeded those in a preceding baseline condition in which no food was delivered. In addition, upon introduction of the delayed reinforcement procedure, correspondence between response patterns and the requirements of the reinforcement schedule increased over baseline levels in the absence of a food contingency. Together with a previous report of Lattal and Gleeson (1990), the present results suggest that response acquisition with delayed reinforcement is a robust phenomenon that may not depend on a mechanically defined response or an immediate external stimulus change to mediate the temporal gap between response and reinforcer.

Animals↗

Choice with certain and uncertain reinforcers in an adjusting-delay procedure.

A discrete-trials adjusting-delay procedure was used to investigate the conditions under which pigeons might show a preference for partial reinforcement over 100% reinforcement, an effect reported in a number of previous experiments. A peck on a red key always led to a delay with red houselights and then food. In each condition, the duration of the red-houselight delay was adjusted to estimate an indifference point. In 100% reinforcement conditions, a peck on a green key always led to a delay with green houselights and then food. In partial-reinforcement conditions, a peck on the green key led either to the green houselights and food or to white houselights and no food. In some phases of the experiment, statistically significant preference for partial reinforcement over 100% reinforcement was found, but this effect was observed in only about half of the pigeons. The effect was largely eliminated when variability in the delay stimulus colors was equated for 50% reinforcement conditions and 100% reinforcement conditions. Idiosyncratic preferences for certain colors or for stimulus variability may be at least partially responsible for the effect.

Animals↗

Effects of reinforcement rate and delay on the acquisition of lever pressing by rats.

The acquisition of lever pressing by naive rats, in the absence of shaping, was studied as a function of different rates and unsignaled delays of reinforcement. Groups of 3 rats were each exposed to tandem schedules that differed in either the first or the second component. First-component schedules were either continuous reinforcement or random-interval 15, 30, 60 or 120 s; second-component schedules were fixed-time 0, 1, 3, 6, 12, or 24 s. Rate of responding was low under continuous immediate reinforcement and higher under random-interval 15 s. Random interval 30-s and 60-s schedules produced lower rates that were similar to each other. Random-interval 120 s controlled the lowest rate in the immediate-reinforcement condition. Adding a constant 12-s delay to each of the first-component schedule parameters controlled lower response rates that did not vary systematically with reinforcement rate. The continuous and random-interval 60-s schedules of immediate reinforcement controlled higher global and first-component response rates than did the same schedules combined with longer delays, and first-component rates showed some graded effects of delay duration. In addition, the same schedules controlled higher second-component response rates in combination with a 1-s delay than in combination with longer delays. These results were related to those from previous studies on acquisition with delayed reinforcement as well as to those from similar reinforcement procedures used during steady-state responding.

Animals↗