Rate Of Key Pecking Research Articles

Reinforcer magnitude and rate dependency: evaluation of resistance-to-change mechanisms.

Under many circumstances, reinforcer magnitude appears to modulate the rate-dependent effects of drugs such that when schedules arrange for relatively larger reinforcer magnitudes rate dependency is attenuated compared with behavior maintained by smaller magnitudes. The current literature on resistance to change suggests that increased reinforcer density strengthens operant behavior, and such strengthening effects appear to extend to the temporal control of behavior. As rate dependency may be understood as a loss of temporal control, the effects of reinforcer magnitude on rate dependency may be due to increased resistance to disruption of temporally controlled behavior. In the present experiments, pigeons earned different magnitudes of grain during signaled components of a multiple FI schedule. Three drugs, clonidine, haloperidol, and morphine, were examined. All three decreased overall rates of key pecking; however, only the effects of clonidine were attenuated as reinforcer magnitude increased. An analysis of within-interval performance found rate-dependent effects for clonidine and morphine; however, these effects were not modulated by reinforcer magnitude. In addition, we included prefeeding and extinction conditions, standard tests used to measure resistance to change. In general, rate-decreasing effects of prefeeding and extinction were attenuated by increasing reinforcer magnitudes. Rate-dependent analyses of prefeeding showed rate-dependency following those tests, but in no case were these effects modulated by reinforcer magnitude. The results suggest that a resistance-to-change interpretation of the effects of reinforcer magnitude on rate dependency is not viable.

DEVELOPMENT OF KEY‐PECKING, PAUSE, AND AMBULATION DURING EXTENDED EXPOSURE TO A FIXED‐INTERVAL SCHEDULE OF REINFORCEMENT

Six pigeons key-pecked under a fixed-interval (FI) 3-min schedule of food presentation. Each pigeon was studied for 200 daily sessions with 15 intervals per session (3,000 total food presentations). Analyses included the examination of latency to first peck (pause), mean rate of key pecking, and ambulation. Characterizations of stable performance were assessed across measures of behavior and evaluated using commonly employed stability criteria. Stability of response rate and pause was identified better by assessments that evaluated variability and trend, rather than just variability. Between-subject differences in rate of acquisition and terminal values of steady-state performance of pause were observed, and stable pause durations took longer to develop than did stable key-pecking rates. Relative variability in response rate and pause duration decreased as the means increased. A temporally organized pattern of key-pecking (the so-called FI scallop) developed within 50 sessions of exposure to the schedule. Overall ambulation decreased during the early sessions of exposure and further analyses showed greater rates of ambulation during the pause than after it for 4 of the 6 pigeons. Performance under the FI 3-min schedule developed relatively slowly, and key-pecking, pause, and ambulation developed at different rates.

Biological and Economic Effects on Responding:Rate and Duration of the Pigeon's Key Peck

Pigeons were studied on a two-component multiple schedule in which key pecking was reinforced on a variable interval (VI) 2-min schedule in both components. In separate phases additional food was delivered on a variable-time (VT) 15-s schedule (response independent) or a VI 15-s schedule (response dependent) in one of the components. In addition to rate, duration of key pecks was measured in an attempt to differentiate the biological and economic effects on key pecking. When components alternated frequently (every 10 s), all pigeons key pecked at a much higher rate during the component with the additional food deliveries, whether response dependent or independent. When components alternated infrequently (every 20 min), pigeons key pecked at high rates at points of transition into the component with the additional food deliveries. Rate of key pecking decreased with time spent in the 20-min component when the additional food was response independent whereas rate of pecking remained elevated when the additional food was response dependent. The additional food deliveries, whether response-independent or response-dependent, however, had no consistent effect on the pigeon's key-peck duration. That is, there were no systematic or reliable shifts in peck duration as would be predicted if short-duration pecks were biologically based. Despite the fact that we were unable to “tag” the biological effect in terms of key-peck duration, the finding that the delivery of response-independent food has different, but predictable effects on responding suggests that animal learning principles can be integrated with species-typical, biological considerations without the need to propose constraints that limit general laws of learning.

Within-session delay-of-reinforcement gradients.

Within-session delay-of-reinforcement gradients were generated with pigeons by progressively increasing delays to reinforcement within each session. In Experiment 1, the effects of imposing progressive delays on variable-interval and fixed-interval schedules were investigated while controlling for simultaneous decreases in reinforcer rate across the session via a within-subject yoked-control procedure. Rate of key pecking decreased as a negatively decelerated function of delay of reinforcement within a session. These rate decreases were greater than those during a yoked-interval session in which the rate of immediate reinforcement decreased at the same rate as it did under the progressive-delay procedure. In Experiment 2, delay-of-reinforcement gradients were shallower when the progressive delay intervals were signaled by a blackout than when they were unsignaled. The delay gradients obtained in each experiment were similar to those generated under conditions in which different delays of reinforcement are imposed across blocks of sessions. The present procedure offers a technique for rapidly generating delay-of-reinforcement gradients that might serve as baselines for assessing the effects of other behavioral and pharmacological variables.

Economic and biological influences on key pecking and treadle pressing in pigeons.

Pigeons were studied on a two-component multiple schedule in which the required operant was, in different conditions, biologically relevant (i.e., key pecking) or nonbiologically relevant (i.e., treadle pressing). Responding was reinforced on a variable-interval (VI) 2-min schedule in both components. In separate phases, additional food was delivered on a variable-time (VT) 15-s schedule (response independent) or a VI 15-s schedule (response dependent) in one of the components. The addition of response-independent food had different effects on responding depending on the operant response and on the frequency with which the components alternated. When components alternated frequently (every 10 s), all pigeons keypecked at a much higher rate during the component with the additional food deliveries, whether response dependent or independent. In comparison, treadle pressing was elevated only when the additional food was response dependent; rate of treadling was lower when the additional food was response independent. When components alternated infrequently (every 20 min), pigeons key pecked at high rates at points of transition into the component with the additional food deliveries. Rate of key pecking decreased with time spent in the 20-min component when the additional food was response independent, whereas rate of pecking remained elevated in that component when the additional food was response dependent. Under otherwise identical test conditions, rate of treadle pressing varied only as a function of its relative rate of response-dependent reinforcement. Delivery of response-independent food thus had different, but predictable, effects on responding depending on which operant was being studied, suggesting that animal-learning procedures can be integrated with biological considerations without the need to propose constraints that limit general laws of learning.

Operant discriminative learning and evidence of subtelencephalic plastic changes after long-term detelencephalation in pigeons.

We analyzed operant discrimination in detelencephalated pigeons and neuroanatomical substrates after long-term detelencephalation. In Experiment I, experimental pigeons with massive telencephalic ablation and control pigeons were conditioned to key peck for food. Successive discrimination was made under alternating red (variable-ratio reinforcement) and yellow (extinction) lights in one key of the chamber. These relations were interchanged during reversal discrimination. The sessions were run until steady-state rates were achieved. Experiment II analyzed the morphology of the nucleus rotundus and optic tectum in long-term detelencephalated and control birds, using a Klüver-Barrera staining and image analyzer system. Detelencephalated birds had more training sessions for response shaping and steady-state behavior (p<0.001), higher red key peck rates during discrimination (p<0.01), and reversal discrimination indexes around 0.50. Morphometric analysis revealed a decreased number of neurons and increased vascularity, associated with increases in the perimeter (p<0.001) in the nucleus rotundus. In the optic tectum, increases in the perimeter (p<0.05) associated with disorganization in the layers arrangement were seen. The data indicate that telencephalic systems might have an essential function in reversal operant discrimination learning. The structural characteristics of subtelencephalic systems after long-term detelencephalation evidence plastic changes that might be related to functional mechanisms of learning and neural plasticity in pigeons.

Effects of D-amphetamine in a temporal discrimination procedure: selective changes in timing or rate dependency?

Two experiments evaluated rate dependency and a neuropharmacological model of timing as explanations of the effects of amphetamine on behavior under discriminative control by time. Four pigeons pecked keys during 60-trial sessions. On each trial, the houselight was lit for a particular duration (5 to 30 s), and then the key was lit for 30 s. In Experiment 1, the key could be lit either green or blue. If the key was lit green and the sample was 30 s, or if the key was lit blue and the sample was 5 s, pecks produced food on a variable-interval 20-s schedule. The rate of key pecking increased as a function of sample duration when the key was green and decreased as a function of sample duration when the key was blue. Acute d-amphetamine (0.1 to 3.0 mg/kg) decreased higher rates of key pecking and increased lower rates of key pecking as predicted by rate dependency, but did not shift the timing functions leftward (toward overestimation) as predicted by the neuropharmacological model. These results were replicated in Experiment 2, in which the key was lit only one color during sessions, indicating that the effects were not likely due to disruption of discriminative control by key color. These results are thus consistent with rate dependency but not with the predictions of the neuropharmacological model.

Pigeons Flexibly Time or Count on Cue

In Experiment 1, pigeons were presented with a sequence of light flashes and cued to peck a key for reward either after a fixed time or after a fixed number of flashes. Curves that showed the rate of key pecking over time within trials indicated that peak rates of response were reached near the fixed time on timing-cued trials and near the fixed number of flashes on counting-cued trials. In Experiment 2, the key cue was shifted from timing to counting or from counting to timing midway through a trial. The peak times reached after the cue change indicated that pigeons kept track of time while cued to count but did not count while cued to time. These findings suggest a basic asymmetry in the dual-mode model of timing and counting.

The effects of morphine on clocked fixed-interval performance: stimulus function or strength of stimulus control?

The present experiment was conducted to determine whether the modification of the effects of morphine by food deprivation would occur under conditions of strong stimulus control. Four pigeons pecked response keys under a multiple fixed-interval (FI) 5-min and clocked fixed-interval (CFI) 5-min schedule of food presentation. During the FI component, the key was lit green and the house light was on throughout. During the CFI component, the key light and house light settings changed in a regular manner as the interval elapsed. In all conditions, pigeons received a range of doses of morphine (1.0-10.0 mg/kg) and saline. In the first condition, pigeons were maintained at 80% of their free-feeding weights. When calculated over entire 5-min intervals, morphine produced dose-dependent decreases in the index of curvature (a measure of temporal patterning) during the FI component, but did not affect the index of curvature during the CFI component. A more detailed analysis revealed that the index of curvature during the last stimulus in the CFI sequence was reduced in a dose-dependent manner, similar to the effects obtained for the entire FI component. When the pigeons were maintained at 70% and 90% of their free-feeding weights, measures of the effects of morphine on temporal patterning did not differ as a function of body weight. The effects of morphine on rates of key-pecking, however, were related to body weight. For three of four pigeons, rates of key-pecking were not suppressed until higher doses, and increases in rates of key-pecking were more common at lower doses, when the pigeons were maintained at 70% of their free-feeding weights, than when they were maintained at 90% of their free-feeding weights. It is suggested that the effects of morphine on CFI performance can be understood in terms of the function of the stimuli that comprise the schedule, rather than the strength of the stimulus control engendered.

The rate-decreasing effects of fentanyl derivatives in pigeons before, during and after chronic morphine treatment.

Mirfentanil is a fentanyl derivative with non-opioid actions, including non-opioid antinociceptive effects in rhesus monkeys. The current study examined the rate-altering effects of mirfentanil and several other compounds in pigeons to assess: 1) the opioid and non-opioid actions of acutely-administered fentanyl derivatives; and 2) the development of cross-tolerance between each of these compounds and morphine. Seven pigeons responded under a fixed-ratio 20 (FR20) schedule of food delivery. In untreated pigeons, fentanyl, morphine, naltrexone, ketamine and three fentanyl derivatives (mirfentanil, OHM3463 and OHM3295) decreased rates of key pecking in a dose-related manner. Naltrexone (0.1-1.0 mg/kg) attenuated the effects of OHM3463 and not mirfentanil or OHM3295, suggesting non-opioid mediation of the rate-decreasing effects for the latter two fentanyl derivatives. Subjects were treated daily with morphine for 9 weeks, up to a dose of 100 mg/kg per day, during which time the dose-effect curves for morphine, fentanyl and OHM3463 shifted rightward 6-, 10- and 2-fold, respectively, indicating the development of tolerance to morphine and cross-tolerance to fentanyl and OHM3463. Dose-effect curves for ketamine, OHM3295 and mirfentanil were not shifted to the right during morphine treatment, and the dose-effect curve for naltrexone was shifted leftward 180-fold. To the extent that rate-decreasing effects are predictive of antinociceptive effects, these data suggest that some fentanyl derivatives might be useful therapeutics under conditions where tolerance develops to morphine-like opioids.

Relations between dose magnitude, subject sensitivity, and the development of tolerance to cocaine-induced behavioral disruptions in pigeons

Keypecking by 12 pigeons, maintained by a fixed-ratio 30 schedule of food presentation, was decreased in rate by acute pre-session injections of cocaine in a dose-dependent manner, with larger doses producing more disruption. A constant dose of cocaine was then injected prior to every session for 40 days. Some subjects received a relatively small dose, some received a medium-sized dose, and others received a large dose. Subsequently, dose-effects were reassessed via once-weekly probe injections, with every other session continuing to be preceded by injection of the daily dose of cocaine. Then a different dose of cocaine was administered daily for 40 more days, after which the dose-effect function was redetermined in like manner. In general, tolerance to cocaine-induced response-rate reductions was most likely to develop when (a) the repeatedly-administered dose of cocaine was relatively small (even without acute effect on keypecking) and (b) the subject's keypecking was disrupted by smaller doses of cocaine in the initial dose-effect assessment. Tolerance was generally observed as a shift in the dose-effect function that, in several cases, could be eliminated by increasing the magnitude of the daily administered dose. In addition, every subject's rate of keypecking following saline injections was lowered after daily exposure to cocaine. These results (a) are partially consistent with the reinforcement-loss account of tolerance to cocaine-induced behavioral disruptions, and (b) support previous observations of withdrawal symptoms following cessation of extended exposure to cocaine.

CHANGEOVER BEHAVIOR AND PREFERENCE IN CONCURRENT SCHEDULES

Pigeons were trained on a multiple schedule of reinforcement in which separate concurrent schedules occurred in each of two components. Key pecking was reinforced with milo. During one component, a variable-interval 40-s schedule was concurrent with a variable-interval 20-s schedule; during the other component, a variable-interval 40-s schedule was concurrent with a variable-interval 80-s schedule. During probe tests, the stimuli correlated with the two variable-interval 40-s schedules were presented simultaneously to assess preference, measured by the relative response rates to the two stimuli. In Experiment 1, the concurrently available variable-interval 20-s schedule operated normally; that is, reinforcer availability was not signaled. Following this baseline training, relative response rate during the probes favored the variable-interval 40-s alternative that had been paired with the lower valued schedule (i.e., with the variable-interval 80-s schedule). In Experiment 2, a signal for reinforcer availability was added to the high-value alternative (i.e., to the variable-interval 20-s schedule), thus reducing the rate of key pecking maintained by that schedule but leaving the reinforcement rate unchanged. Following that baseline training, relative response rates during probes favored the variable-interval 40-s alternative that had been paired with the higher valued schedule. The reversal in the pattern of preference implies that the pattern of changeover behavior established during training, and not reinforcement rate, determined the preference patterns obtained on the probe tests.

Opioid-induced response-rate decrements in pigeons responding under variable-interval schedules

Opioid drugs may produce some of their behavioral effects by altering the effectiveness of reinforcing events. The present investigation examined effects of two opioids (methadone and buprenorphine) on pigeons' key-pecking. Different reinforcement rates were arranged using five variable-interval (VI) food-presentation schedules, permitting an application of Heyman's Matching Law analysis and Nevin's Resistance to Change Hypothesis to behavioral actions of opioid drugs. Key-pecking by four pigeons was reinforced by 3-s access to mixed grain under a five component multiple VI schedule. VI values used were 5, 10, 30, 75 and 150s. Each component was in effect for 5min followed by 1min of darkness. Peck rates were high under the VI5-s and VI10-s schedules. As the mean interval value increased, peck rates decreased. Methadone (0.5, 1.5, 2.5, 3.75, and 5.0mg/kg) and buprenorphine (0.25, 0.5, 1.0, 3.0, and 5.0mg/kg), administered 30min prior to experimental sessions, dose dependently decreased peck rates in all subjects. Peck rates under longer VI schedules (75 and 150s) tended to be more greatly disrupted than those maintained under shorter VI schedules (e.g. 10s). Dose-by-dose analysis of key-pecking rate changes under each schedule, and analysis of drug-induced changes in Matching Law parameters suggest that peck rate decreases resulted, in part, from decrements in the reinforcer's ability to maintain behavior.

Selective sensitivity of schedule-induced activity to an operant suppression contingency.

The sensitivity of pigeons' schedule-induced activity to operant consequences was studied in two experiments. During a 30-s interval between food presentations, a keylight stimulus brightened incrementally. Stable terminal key pecking and interim locomotor activity developed. An operant "setback" contingency was applied to activity. The contingency arranged for locomotor movements (detected by a nine-panel floorboard) to be followed by a resetting of the keylight brightness to a dimmer value and a 1-s delay of reinforcement (for individual responses). Experiment 1 showed that activity patterns were highly sensitive to their operant consequences. Accompanying key-peck rates were only transiently affected. In Experiment 2, the setback contingency was imposed during restricted portions of the trial, and differential operant control of activity was demonstrated. However, birds in this study produced higher rates of key pecking as activity rates were reduced. These results suggest that although schedule-induced activity arises in response to the temporal arrangement of stimulus events, this behavior may retain considerable sensitivity to response-consequence relations.

Key pecking of pigeons under variable-interval schedules of briefly signaled delayed reinforcement: effects of variable-interval value.

Key pecking of 4 pigeons was maintained under a multiple variable-interval 20-s variable-interval 120-s schedule of food reinforcement. When rates of key pecking were stable, a 5-s unsignaled, nonresetting delay to reinforcement separated the first peck after an interval elapsed from reinforcement in both components. Rates of pecking decreased substantially in both components. When rates were stable, the situation was changed such that the peck that began the 5-s delay also changed the color of the keylight for 0.5 s (i.e., the delay was briefly signaled). Rates increased to near-immediate reinforcement levels. In subsequent conditions, delays of 10 and 20 s, still briefly signaled, were tested. Although rates of key pecking during the component with the variable-interval 120-s schedule did not change appreciably across conditions, rates during the variable-interval 20-s component decreased greatly in 1 pigeon at the 10-s delay and decreased in all pigeons at the 20-s delay. In a control condition, the variable-interval 20-s schedule with 20-s delays was changed to a variable-interval 35-s schedule with 5-s delays, thus equating nominal rates of reinforcement. Rates of pecking increased to baseline levels. Rates of pecking, then, depended on the value of the briefly signaled delay relative to the programmed interfood times, rather than on the absolute delay value. These results are discussed in terms of similar findings in the literature on conditioned reinforcement, delayed matching to sample, and classical conditioning.

Generalization peak shift for autoshaped and operant key pecks.

Pigeons acquired discriminated key pecking between 528- and 540-nm stimuli by either a response-reinforcer (operant group) or a stimulus-reinforcer (autoshaped group) contingency, with other training-schedule parameters comparable over groups. For the birds in the operant group, key pecks intermittently produced grain in the presence of one hue on the key (positive stimulus) but not in the other (negative stimulus). For the birds in the autoshaped group, pecking emerged when grain was intermittently presented independently of key pecking during one key color but was not presented during the other key color. Two independent contingency assays, peck-location comparisons and elimination of differences in reinforcement rate, confirmed the effectiveness of the two training procedures in establishing operant or respondent control of key pecking. After reaching a 10:1, or better, discrimination ratio between key pecks during the two key colors, the birds received a wavelength generalization test. Criterion baseline key-peck rates were comparable for operant and autoshaped groups prior to testing. On the generalization test, performed in extinction, all birds pecked most at a stimulus removed from the positive training stimulus in the direction away from the negative stimulus. In testing, autoshaped "peak" rates (24.5 to 64.9 pecks per minute) were from 33% to 80% higher than rates in the presence of the training stimuli. Respondent peak shift rarely has been reported heretofore, and never this consistently and robustly. These results further confirm the similarity of perceptual processing in classical and operant learning. They are discussed in terms of Spence's gradient-interaction theory and Weiss' (1978) two-process model of stimulus control.

Novelty versus retrieval cue value in the study of long-term memory in pigeons.

Six experiments investigated whether a training context distinguished by an ambient odor (isoamyl acetate or eucalyptus oil) might serve as a retrieval cue for pigeons by virtue of its mere presence during reinforced learning. The rate of key pecking in the presence of 1 of 2 odors that has accompanied reinforced responding, relative to the rate in the alternative (novel) odor (Experiments 2 and 4), was compared with the relative response rate in the presence of an odor that was familiar but had not accompanied reinforced responding (Experiment 1 and 3)

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.

Effects on opioid-induced rate reductions by doxepin and bupropion

Twelve pigeons key-pecked under a multiple variable interval 15-second, variable interval 150-second schedule of food reinforcement. Effects of two opioid drugs, buprenorphine and methadone, were determined alone and in combination with chronic daily administration of the antidepressants doxepin or bupropion. Methadone initially produced dose-dependent key-pecking rate reductions when administered acutely, prior to the session, while buprenorphine produced key-pecking rates that reached a plateau at 50–80% of baseline rate and were not reduced further by higher doses. Neither doxepin nor bupropion, given alone, had lasting effects on key-pecking rates. Chronic daily doxepin administration significantly attenuated methadone-induced response rate reductions. Bupropion reduced the effect of the highest methadone dose, but this effect was mitigated by the development of opioid tolerance. Unlike bupropion, doxepin interfered with the development of opioid tolerance. Neither antidepressant systematically altered effects of buprenorphine on key-pecking.

Effects of trial duration on overall and momentary rates of maintained autoshaped keypecking: Choice and single-stimulus trials

Pigeons were exposed to differentially cued autoshaping trials in which conditioned stimuli were followed by food after 6 or 14 sec. Average and momentary rates of keypecking were examined on two types of unreinforced test trials: single-stimulus probe trials and simultaneous choice trials, each 40 sec in duration. Rates averaged over the 40-sec test trials did not favor the cue associated with the shorter delay to food (the short-delay cue) on either type of test trial; however, average rates prior to the scheduled time of food delivery were reliably higher for the short-delay cue on choice trials. Momentary rates of keypecking during choice trials varied as a function of both cue and elapsed time from trial onset. At short elapsed trial times, rate of pecking was higher for the short-delay cue, with this difference reversing at longer times. A reversal of the programmed relation between key color and delay to food presentation for 5 birds confirmed the generality of these findings. Implications of these data for models of Pavlovian conditioning and for methods of assessing conditioned response strength are discussed.