Biasing the pacemaker in the behavioral theory of timing

In the behavioral theory of timing, pacemaker rate is determined by overall rate of reinforcement. A two-alternative free-operant psychophysical procedure was employed to investigate whether pacemaker period was also sensitive to the differential rate of reinforcement. Responding on a left key during the first 25 s and on a right key during the second 25 s of a 50-s trial was reinforced at variable intervals, and variable-interval schedule values during the two halves of the trials were varied systematically. Responding on the right key during the first 25 s and on the left key during the second 25 s was not reinforced. Estimates of pacemaker period were derived from fits of a function predicted by the behavioral theory of timing to right-key response proportions in consecutive 5-s bins of the 50-s trial. Estimates of pacemaker period were shortest when the differential reinforcer rate most strongly favored right-key responses, and were longest when the differential reinforcer rate most strongly favored left-key responses. The results were consistent with the conclusion that pacemaker rate is influenced by relative reinforcer rate.     

Rapid determinations of preference in multiple concurrent-chain schedules.

With concurrent chains arranged for a pigeon's key pecks, pecks on two concurrently available initial-link keys (left and right) respectively produce separately operating terminal links (A and B). Preferences for terminal link A over terminal link B are usually calculated as deviations of relative initial-link response rates (left divided by total pecks) from those during baseline conditions, when A equals B. Baseline preferences, however, are often variable and typically are determined indirectly (e.g., with unequal A and B, reversing left-right assignments of A and B over sessions and estimating the baseline from differences between the relative rates generated). Multiple concurrent-chain schedules, with components each consisting of a pair of concurrent chains, speed the determination of preferences by arranging A and B and their reversal within sessions. In two experiments illustrating the feasibility of this procedure, one component operated with circles projected on initial-link keys and the other with pluses; when left and right initial-link pecks respectively produced terminal links A and B in one component, they produced B and A in the other. Even as the baselines fluctuated, preference was observable within sessions as the difference between relative initial-link response rates in the two components. The first experiment demonstrated the rapid development of preferences when terminal links A and B consisted of fixed-interval 15-s and 30-s schedules. The second demonstrated the sensitivity of the procedure to preference for a fixed-interval 30-s schedule operating for pecks on either of two keys (free choice) over its operating for pecks on only a single key (forced choice).     

Food-deprivation level alters the effects of morphine on pigeons'' key pecking.

Four pigeons pecked response keys under a multiple fixed-ratio 30 fixed-interval 5-min schedule of food presentation. Components alternated separated by 15-s timeouts; each was presented six times. Pigeons were maintained at 70%, 85%, and greater than 90% of their free-feeding weights across experimental conditions. When response rates were stable, the effects of morphine (0.56 to 10.0 mg/kg) and saline were investigated. Morphine reduced response rates in a dose-dependent manner under the fixed-ratio schedule and at high doses under the fixed-interval schedule. In some cases, low doses of morphine increased rates under the fixed-interval schedule. When pigeons were less food deprived, reductions in pecking rates occurred at lower doses under both schedules for 3 of 4 birds compared to when they were more food deprived. When pigeons were more food deprived, low doses of morphine increased rates of pecking in the initial portions of fixed intervals by a greater magnitude. Thus, food-deprivation levels altered both the rate-decreasing and rate-increasing effects of morphine. These effects may share a common mechanism with increased locomotor activity produced by drugs and with increased drug self-administration under conditions of more severe food deprivation.     

Pigeons presented with sequences of light flashes use behavior to count but not to time

On randomly ordered trials, pigeons were presented with either a blue or a white key that flashed red for 200 ms at a fast (2 flashes/s), medium (1 flash/s), or slow (0.5 flashes/s) rate. The blue key signaled a fixed-interval (FI) schedule in which the first response after 20 s was reinforced, and the white key signaled a fixed-number (FN) schedule in which the first response after 20 flashes was reinforced. In Experiments 1 and 2, pigeons showed depressed responding to the flash on FI-cued trials and accelerated responding to the flash on FN-cued trials. When the response key was periodically blacked out in Experiments 3 and 4, counting but not timing was eliminated.     

Concurrent performances: inhibition of one response by reinforcement of another

In an analysis of interactions between concurrent performances, variable-interval reinforcement was scheduled, in various sequences, for both keys, for only one key, or for neither key of a two-key pigeon chamber. With changeover delays of 0.5 or 1.0 sec, and with each key's reinforcements discriminated on the basis of key-correlated feeder stimuli, reinforcement of pecks on one key reduced the pecking maintained by reinforcement on the other key. The decrease in pecking early after reinforcement was discontinued on one key was not substantially affected by whether pecks on the other key were reinforced, but after reinforcement was discontinued on both keys, reinstatement of reinforcement for one key sometimes produced transient increases in pecking on the other key. Correlating the availability of right-key reinforcements with a stimulus, which maintained right-key reinforcement while reducing right-key pecking to negligible levels, demonstrated that these interactions depended on concurrent reinforcement, not concurrent responding. Thus, reinforcement of a response, but not necessarily the occurrence of the response, inhibits other reinforced responses. Compared with accounts in terms of excitatory effects of extinction, often invoked in treatments of behavioral contrast, this inhibitory account has the advantage of dealing only with observed dimensions of behavior.     

Pigeons may not remember the stimuli that reinforced their recent behavior

In two experiments the conditioned reinforcing and delayed discriminative stimulus functions of stimuli that signal delays to reinforcement were studied. Pigeons' pecks to a center key produced delayed-matching-to-sample trials according to a variable-interval 60-s (or 30-s in 1 pigeon) schedule (Experiment 1) or a multiple variable-interval 20-s variable-interval 120-s schedule (Experiment 2). The trials consisted of a 2-s illumination of one of two sample key colors followed by delays ranging across phases from 0.1 to 27.0 s followed in turn by the presentation of matching and nonmatching comparison stimuli on the side keys. Pecks to the key color that matched the sample were reinforced with 4-s access to grain. Under some conditions of Experiment 1, pecks to nonmatching comparison stimuli produced a 4-s blackout and the start of the next interval. Under other conditions of Experiment 1 and each condition of Experiment 2, pecks to nonmatching stimuli had no effect and trials ended only when pigeons pecked the other, matching stimulus and received food. The functions relating pretrial response rates to delays differed markedly from those relating matching-to-sample accuracy to delays. Specifically, response rates remained relatively high until the longest delays (15.0 to 27.0 s) were arranged, at which point they fell to low levels. Matching accuracy was high at short delays, but fell to chance at delays between 3.0 and 9.0 s. In Experiment 2, both matching accuracy and response rates remained high over a wider range of delays in the variable-interval 120-s component relative to the variable-interval 20-s component. The difference in matching accuracy between the components was not due to an increased tendency in the variable-interval 20-s component toward proactive interference following short intervals. Thus, under these experimental conditions the conditioned reinforcing and the delayed discriminative functions of the sample stimulus depended on the same variables (delay and variable-interval value), but were nevertheless dissociated.     

Drug discrimination under a concurrent fixed-interval fixed-interval schedule.

Pigeons were trained to discriminate 5.0 mg/kg pentobarbital from saline under a concurrent fixed-interval (FI) FI schedule of food presentation on which, after pentobarbital administration, responses on one key were reinforced with food under an FI 60-s component and responses on the other key were reinforced under an FI 240-s component. After saline administration, the schedule contingencies on the two keys were reversed. After both pentobarbital and saline, pigeons responded more frequently on the key on which responses had been programmed to produce the reinforcer under the FI 60 component of the concurrent schedule. The schedule was changed to concurrent FI 150 FI 150 s for drug-substitution tests. In each bird, increasing doses of pentobarbital, ethanol, and chlordiazepoxide produced increases in the proportion of responses on the key on which responses had been reinforced under the FI 60 component after pentobarbital administration during training sessions. The proportion of responses on that key was slightly lower for ethanol than for chlordiazepoxide and pentobarbital. At a dose of pentobarbital higher than the training dose, responding decreased on the key that had been reinforced under the FI 60 component during training sessions. Phencyclidine produced less responding on the key programmed under the FI 60-s component than did pentobarbital. Methamphetamine produced responding primarily on the key on which responses had been reinforced under the FI 60-s component after saline administration.     

The behavioral theory of timing: Reinforcer rate determines pacemaker rate

In the behavioral theory of timing, pulses from a hypothetical Poisson pacemaker produce transitions between states that are correlated with adjunctive behavior. The adjunctive behavior serves as a discriminative stimulus for temporal discriminations. The present experiments tested the assumption that the average interpulse time of the pacemaker is proportional to interreinforcer interval. Responses on a left key were reinforced at variable intervals for the first 25 s since the beginning of a 50-s trial, and right-key responses were reinforced at variable intervals during the second 25 s. Psychometric functions relating proportion of right-key responses to time since trial onset, in 5-s intervals across the 50-s trial, were sigmoidal in form. Average interpulse times derived by fitting quantitative predictions from the behavioral theory of timing to obtained psychometric functions decreased when the interreinforcer interval was decreased and increased when the interreinforcer interval was increased, as predicted by the theory. In a second experiment, average interpulse times estimated from trials without reinforcement followed global changes in interreinforcer interval, as predicted by the theory. Changes in temporal discrimination as a function of interreinforcer interval were therefore not influenced by the discrimination of reinforcer occurrence. The present data support the assumption of the behavioral theory of timing that interpulse time is determined by interreinforcer interval.     

Conjunctive schedules of reinforcement: I. Rate-dependent effects of pentobarbital and d-amphetamine

Key pecking of pigeons was maintained under conjunctive schedules of food presentation in which both a fixed-interval and a fixed-ratio schedule had to be completed before a peck produced food. For two pigeons, pecks on a single key completed both schedule requirements (fixed-interval 3-min, fixed-ratio 50 for one bird, fixed-interval 5-min, fixed-ratio 50 for the second). For two other pigeons, each requirement was scheduled on a separate key. On the two-key schedule, a peck after 5 min on the key scheduling the fixed-interval requirement produced food if at least 10 pecks had occurred on the ratio key (conjunctive fixed-interval 5-min, fixed-ratio 10). When each requirement was scheduled on a separate key, response rates on the fixed-ratio key were generally higher in the early portion of the interval and declined as the interval progressed; responding on the fixed-interval key, once initiated, typically remained at a constant rate throughout the interval. Responding under the single-key schedule was characterized by a high rate early in the interval; this then changed to a lower rate that continued until a peck produced food. For all pigeons, increases in response rates with pentobarbital and d-amphetamine were inversely related to the control rate of responding. When equivalent rates on each key of the two-key schedule were compared, both drugs increased rates on the fixed-ratio key less. Although the effects of both drugs were rate dependent, each drug differentially modified the pattern of responding under the single-key schedule.     

Chained and tandem fixed-interval schedules of punishment

Pigeons' key pecks were both punished with electric shock on four-component chained and tandem fixed-interval schedules and reinforced on a variable-interval schedule of food presentation. Pecking was suppressed less in the early components of the chained schedule than in the early components of the tandem schedule. Related multiple and mixed schedules of punishment were also presented; these schedules were identical to the chained and tandem schedules, respectively, except that components changed independently of responding. Similar effects were obtained, in that responding was suppressed in all components of the mixed schedule and only in the fourth component of the multiple schedule of punishment. The performances maintained on the chained and tandem schedules of punishment were generally symmetrical to those found in analogous chained and tandem schedules of food reinforcement.     

Timing light and tone signals in pigeons

Pigeons' ability to time light and tone stimuli was examined in four experiments. In Experiment 1, two groups of pigeons were trained to discriminate between 2- and 8-s durations of lights or tones and then were transferred to reversal or nonreversal discriminations in the alternate modality. Pigeons learned the light discrimination faster than the tone discrimination and showed immediate positive intermodal transfer from tone to light but not from light to tone. In Experiments 2-4, the peak procedure was used to study birds' timing of 15- and 30-s fixed-interval light and tone signals. Peak times on empty trials under baseline conditions closely approximated the length of fixed-interval signals. When pigeons were tested with time-outs and intermodal switches introduced midway through an empty trial, they tended to reset the timing mechanism and begin timing again from 0 s. With both estimation and production procedures, pigeons were less accurate when timing the tone stimuli than when timing the light stimuli. A comparison of these data with data from timing experiments with rats suggests several possible differences in timing processes between pigeons and rats.     

Remote effects of aversive contingencies: Disruption of appetitive behavior by adjacent avoidance sessions

Disruption of ongoing appetitive behavior before and after daily avoidance sessions was examined. After baselines of appetitive responding were established under a fixed-interval 180-s schedule of food presentation, 4 rats were exposed to 40-min sessions of the appetitive schedule just prior to 100-min sessions of electric shock postponement, while another 4 rats received the 40-min appetitive sessions just following daily sessions of shock postponement. In all 8 subjects, fixed-interval response rates decreased relative to baseline levels, the effect being somewhat more pronounced when the avoidance sessions immediately followed. The disruption of fixed-interval responding was only partially reversed when avoidance sessions were discontinued. During the initial exposure to the avoidance sessions, patterns of responding under the fixed-interval schedule were differentially sensitive to disruption, with high baseline response rates generally more disturbed than low rates. These disruptions were not systematically related to changes in reinforcement frequency, which remained fairly high and invariant across all conditions of the experiment; they were also not systematically related to the response rates or to the shock rates of the adjacent avoidance sessions. The results, while qualitatively resembling patterns of conditioned suppression as typically studied, occurred on a greatly expanded time scale. As disruption of behavior extending over time, the present data suggest that some forms of conditioned suppression are perhaps best viewed within a larger temporal context.     

Control of key pecking by the duration of a visual stimulus

Pigeons' key pecks were brought under the control of the duration of a visual stimulus in one-key and two-key procedures. In the one-key procedure, pecks were reinforced after presentations of a long-duration stimulus but not after presentations of a short-duration stimulus. In the two-key procedure, left-key pecks were reinforced after the long-duration stimulus and right-key pecks after the short-duration stimulus. In both procedures, the long-duration stimulus was 10 sec, and the short-duration stimulus was increased from 1 to 8 sec in 1-sec steps. Discriminative control developed with both procedures, but with greater accuracy in the two-key procedure, in which a difference threshold was obtained at short-duration values between 7 and 8 sec, or about 2.5 sec shorter than the long-duration stimulus.     

Cocaine's effects on food-reinforced pecking in pigeons depend on food-deprivation level.

Four pigeons deprived to 80% of their laboratory free-feeding weights pecked keys under a multiple fixed-ratio 30 fixed-interval 5-min schedule of food presentation. Components alternated strictly with 15-s timeouts separating them; each was presented six times. When rates of pecking were stable, 2 pigeons' weights were reduced to 70%, and the other 2 pigeons' weights were increased to 82.5% to 85% of free-feeding levels. Cocaine (1.0, 3.0, 5.6, and 10.0 mg/kg and saline) was administered 5 min prior to sessions. When each dose had been tested twice, pigeons' weights were adjusted to the level that they had not yet experienced, and cocaine was tested again. Cocaine reduced response rates in a dose-dependent manner under the fixed-ratio schedule and under the fixed-interval schedule at high doses, and increased rates under the fixed-interval schedule at low low doses. Reductions in pecking rates occurred at lower doses under both schedules in 3 of 4 pigeons when they were less food deprived compared to when they were more food deprived. Low doses of cocaine increased low baseline rates of pecking in the initial portions of the fixed-interval schedules by a greater magnitude when pigeons were more food deprived. Thus, food-deprivation levels altered both the rate-decreasing and rate-increasing effects of cocaine. The implications of these results for the mechanisms by which food deprivation increases cocaine self-administration and for the dependence of cocaine's effects on the baseline strength of operant behavior are discussed.     

A pharmacological examination of the resistance-to-change hypothesis of response strength.

The effects of d-amphetamine sulfate, sodium pentobarbital, haloperidol, and cholecystokinin-octapeptide were examined within the context of Nevin's (1974, 1979) resistance-to-change hypothesis of response strength. In three experiments, rats' responding was reinforced by delivery of food under chained random-interval 30-s random-interval 30-s, multiple fixed-interval 30-s fixed-interval 120-s, or multiple random-interval 30-s random-interval 120-s schedules. Each rat received several doses of each drug and changes in response rate were measured. The resistance-to-change hypothesis predicts greater disruption of response rate relative to baseline in the initial component of the chained schedule and in the 120-s component of the multiple schedules. In the chained schedule cholecystokinin-octapeptide produced greater reductions in response rate relative to baseline in the initial component. However, no differences between components were observed with haloperidol or sodium pentobarbital, and high doses of d-amphetamine reduced response rate in the terminal component relatively more than in the initial component. In the multiple schedules either no differences were observed between components or response rate was reduced more relative to baseline in the 30-s component. The data fail to support the notion that drugs may be viewed within the same context as other response disruptors such as extinction, satiation, and the presentation of alternative reinforcement.     

Effects of reinforcement history on response rate and response pattern in periodic reinforcement

Several researchers have suggested that conditioning history may have long-term effects on fixed-interval performances of rats. To test this idea and to identify possible factors involved in temporal control development, groups of rats initially were exposed to different reinforcement schedules: continuous, fixed-time, and random-interval. Afterwards, half of the rats in each group were studied on a fixed-interval 30-s schedule of reinforcement and the other half on a fixed-interval 90-s schedule of reinforcement. No evidence of long-term effects attributable to conditioning history on either response output or response patterning was found; history effects were transitory. Different tendencies in trajectory across sessions were observed for measures of early and late responding within the interreinforcer interval, suggesting that temporal control is the result of two separate processes: one involved in response output and the other in time allocation of responding and not responding.     

Studies of operant and reflexive key pecks in the pigeon

The duration of pigeons' key pecks was studied in three experiments. Experiment I revealed that key pecks early in exposure to continuous reinforcement were of short duration, as were key pecks observed on an omission procedure in which pecks prevented food delivery. Key pecks later in exposure to continuous reinforcement, and those that occurred on positive automaintenance procedures, were of long duration. In Experiment II, pigeons were exposed to fixed-interval and fixed-ratio reinforcement schedules, and durations were recorded separately for each quarter of each interval or ratio. On fixed interval, durations were shorter in the first quarter of each interval than in subsequent quarters; on fixed ratio, durations were longer in the first quarter of the ratio than in subsequent quarters. These data parallel observations of concurrent operant responding and salivation in dogs. In Experiment III, pigeons were exposed to a discrete trial, differential-reinforcement-of-low-rate 6-sec schedule. Durations of responses in the first 2 sec of the trial were substantially shorter than those of responses that occurred later. The data from all three experiments support the view that the pigeon's "key peck" actually consists of two subclasses of peck, one reflexive and one operant.     

Decision rules and signal detectability in a reinforcement-density discrimination

Two probabilistic schedules of reinforcement, one richer in reinforcement, the other leaner, were overlapping stimuli to be discriminated in a choice situation. One of two schedules was in effect for 12 seconds. Then, during a 6-second choice period, the first left-key peck was reinforced if the richer schedule had been in effect, and the first right-key peck was reinforced if the leaner schedule had been in effect. The two schedule stimuli may be viewed as two binomial distributions of the number of reinforcement opportunities. Each schedule yielded different frequencies of 16 substimuli. Each substimulus had a particular type of outcome pattern for the 12 seconds during which a schedule was in effect, and consisted of four consecutive light-cued 3-second T-cycles, each having 0 or 1 reinforced center-key pecks. Substimuli therefore contained 0 to 4 reinforcers. On any 3-second cycle, the first center-key peck darkened that key and was reinforced with probability .75 or .25 in the richer or leaner schedules, respectively. In terms of the theory of signal detection, detectability neared the maximum possible d′ for all four pigeons. Left-key peck probability increased when number of reinforcers in a substimulus increased, when these occurred closer to choice, or when pellets were larger for correct left-key pecks than for correct right-key pecks. Averaged over different temporal patterns of reinforcement in a substimulus, substimuli with the same number of reinforcers produced choice probabilities that matched relative expected payoff rather than maximized one alternative.     

Behavior-dependent reinforcer-rate changes in concurrent schedules: A further analysis

Six pigeons were trained on concurrent variable-interval schedules in three different procedures. The first procedure was a standard concurrent schedule, and the relative reinforcer frequency for responding was varied. The second was a schedule in which a relative left-key response rate (over a fixed period of time) exceeding .75 produced, in the next identical time period a higher reinforcer rate on the right key. If this criterion was not exceeded, equal reinforcer rates were arranged on the two keys in this period. This was the dependent procedure. In the third (independent) procedure, the periods of higher right-key reinforcer rates occurred with the same probability as in the second procedure, but occurred independently of behavior. In the second and third procedures, the fixed-time period (window) was varied from 5 s to 60 s, and to 240 s in the second procedure only. Performance on the two keys was similar in the concurrent and independent procedures. The procedure used in the dependent conditions generally affected performance when the windows were shorter than about 30 s. Models of performance that assume that subjects do not discriminate changes in local relative reinforcer rates cannot account for the data. Moreover, existing models are inherently unable to account for the effects of contingencies of reinforcement between responding on one alternative and gaining reinforcers on another that are arranged or that emerge as a result of time allocated to alternative schedules. Undermatching on concurrent variable-interval schedules may result from such emergent contingencies.     

Timing, remembering, and discrimination

Four pigeons were first trained in a timing procedure. In one condition, each trial began with the presentation of an X on the center key, followed by a delay (short or long), after which two side keys were lit. If the delay was short, pecks to the red side key were reinforced. If the delay was long, pecks to the green side key were reinforced. In a second condition, the opposite contingencies applied following presentation of a square on the center key. Choice responses were then tested at 10 time intervals ranging from short to long (1 to 4 s and 4 to 7 s in different conditions). The two timing conditions were combined to create a remembering condition in which correct responding depended upon discrimination of both the sample stimulus (X or square) and the delay interval (short or long). Choices varied systematically across delay in timing conditions, but in remembering conditions, accurate choice at the training delays did not initially generalize to intermediate delays. However, with prolonged training in the remembering task, the response pattern began to resemble that of the timing conditions. Generalization gradients were asymmetrical, in accordance with Weber's Law, in that greater generalization occurred with longer delays than with shorter delays.