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.     

Interval and ratio reinforcement of a complex sequential operant in pigeons

Pigeons were required to produce exactly four pecks on each of two keys in any order for reinforcement. Correct response sequences were reinforced on either fixed-interval two-minute or fixed-ratio four schedules, with each correct sequence treated as a single response. Each pigeon developed a particular dominant sequence that accounted for more than 80% of all sequences. Sequence stereotypy was relatively unaffected by the temporal properties of the fixed-interval and fixed-ratio schedules. Response time (time from the first response in each sequence to the last) was also relatively unaffected by the temporal properties of the schedules. In contrast, response latency (time from end of one sequence to the beginning of the next) was markedly affected by the schedules. Latencies were long early in the interreinforcement interval and got shorter as the interreinforcement interval progressed. These data suggest that stereotyped response sequences become functional behavioral units, resistant to disruption or alteration by reinforcement variables that ordinarily influence the temporal spacing of individual responses.     

Cocaine and food deprivation: effects on food-reinforced fixed-ratio performance in pigeons.

Key pecking by 6 pigeons was maintained by a fixed-ratio 30 schedule of food presentation while body weights were 80% of free-feeding weights. Acute administration of cocaine (0.3 to 13.0 mg/kg, i.m.) dose-dependently decreased response rates. Dose-effect curves were shifted to the right when 3 of the 6 pigeons were maintained at 70% of free-feeding weights and were shifted to the left when the other 3 pigeons were maintained at 90% of free-feeding weights. Then a dose of cocaine that initially decreased response rates by more than 95% of control rates was administered before each daily session. Comparable degrees of tolerance to these rate-decreasing effects developed in the two groups. The rate at which responding recovered was relatively rapid for pigeons in the 70% free-feeding-weight group and was slower for 2 of the 3 pigeons in the 90% free-feeding-weight group. When body weights were then increased from 70% to 80% or were decreased from 90% to 80% of free-feeding weight, performance was disrupted initially only for pigeons whose weight went from 70% to 80% of free feeding. In the present experiment the degree of deprivation may have indirectly influenced the degree of tolerance that developed to cocaine's response rate-decreasing effects because it directly influenced the dose chosen to be administered chronically. The degree of deprivation appeared to have a more direct influence on the rate at which tolerance developed.     

Selective punishment early and late in fixed-ratio schedules of food reinforcement

Pigeons key pecked for grain on a fixed-ratio 100 schedule; electric shocks occurred intermittently at the fifteenth or eighty-fifth response in the ratio. In Experiment I, shock was at the fifteenth response for two birds, and at the eighty-fifth response for two others, in every sixth, twelfth, or eighteenth ratio. Rate of responding decreased as frequency of shock increased, and the pattern of responding included an increased initial pause and low rates or pause-run sequences that extended further into the ratio when shock was at the fifteenth response than when it was at the eighty-fifth response. Shock early in the ratio engendered longer initial pauses than shock late in the ratio. In Experiment II, four birds responded on a two-component multiple schedule in which shock occurred at the fifteenth response of the third ratio in the presence of a white keylight and at the eighty-fifth response of the third ratio in the presence of a green keylight. The overall rates of responding decreased as shock intensity increased. All four birds responded differentially to the white and green keylights, but with a pattern that varied between birds. In general, punishment reduced the probability of responses that preceded it, regardless of the ordinal position of those responses. Both studies confirm that the probability of responding is reduced less by aversive stimuli produced late in a fixed-ratio than by aversive stimuli produced early in a fixed-ratio.     

Neurochemical changes correlated with behavior maintained under fixed-interval and fixed-ratio schedules of reinforcement.

Key pecking of 4 pigeons was maintained under a multiple 3-min fixed-interval, 30-response fixed-ratio schedule of food presentation. Only one schedule was in effect during an experimental session, and each was correlated with a different keylight stimulus and location (left vs. right). The different schedule components alternated across days or weeks. Cerebrospinal fluid was collected from chronically implanted intracerebroventricular cannulae following sessions with the different schedules, as well as following sessions in which reinforcement was withheld (extinction), when response-independent food was delivered, and when the experimental chamber was dark and there were no scheduled events. Metabolites of the neurotransmitters serotonin, norepinephrine, and dopamine were assayed in cerebrospinal fluid using high-performance liquid chromatography with electrochemical detection. Compared to the fixed-ratio condition, responding maintained under the fixed-interval schedule resulted in consistently higher levels of the serotonin metabolite 5-hydroxyindoleacetic acid and of the dopamine metabolite homovanillic acid in all pigeons. Levels of 3-methoxy-4-hydroxyphenylethylene glycol, a metabolite of norepinephrine, and dihydroxyphenylacetic acid, another dopamine metabolite, were also higher in 3 of the 4 pigeons following exposure to the fixed-interval schedules when compared to levels of these metabolites after exposure to the fixed-ratio schedule. Extinction of fixed-ratio responding resulted in large increases in 5-hydroxyindoleacetic acid compared to levels of this metabolite under the fixed-ratio schedule, whereas this serotonin metabolite decreased during extinction of responding under the fixed-interval schedule. Control procedures suggested that the neurochemical changes were not related to the rate of responding but were a function of the specific experimental conditions. Distinctive neurochemical changes that accompany schedule-controlled responding show the sensitivity of the neurochemical environment to behavioral contingencies and demonstrate further the profound impact that such contingencies have on biobehavioral processes.     

Effects of reinforcer duration on responding in two-link chained interval schedules

Each of five pigeons was exposed to two or more durations of access to mixed grains on two-link, chained, interval schedules in which both links were identical fixed-interval or variable-interval schedules. Response rates were an increasing function of reinforcer duration for both initial and terminal links. For both types of interval schedules, initial-link response rates were more sensitive to reinforcer duration than were terminal-link response rates. The present results, together with prior ones, suggest that chaining and choice procedures are each sufficient for demonstrating substantial sensitivity of responding to changes in reinforcer duration.     

Waiting in pigeons: the effects of daily intercalation on temporal discrimination.

Pigeons trained on cyclic-interval schedules adjust their postfood pause from interval to interval within each experimental session. But on regular fixed-interval schedules, many sessions at a given parameter value are usually necessary before the typical fixed-interval "scallop" appears. In the first case, temporal control appears to act from one interfood interval to the next; in the second, it appears to act over hundreds of interfood intervals. The present experiments look at the intermediate case: daily variation in schedule parameters. In Experiments 1 and 2 we show that pauses proportional to interfood interval develop on short-valued response-initiated-delay schedules when parameters are changed daily, that additional experience under this regimen leads to little further improvement, and that pauses usually change as soon as the schedule parameter is changed. Experiment 3 demonstrates identical waiting behavior on fixed-interval and response-initiated-delay schedules when the food delays are short (less than 20 s) and conditions are changed daily. In Experiment 4 we show that daily intercalation prevents temporal control when interfood intervals are longer (25 to 60 s). The results of Experiment 5 suggest that downshifts in interfood interval produce more rapid waiting-time adjustments than upshifts. These and other results suggest that the effects of short interfood intervals seem to be more persistent than those of long intervals.     

Induced attack during fixed-ratio and matched-time schedules of food presentation

Adjunctive or induced behavior is generated during a variety of schedules of reinforcement. Several theoretical conceptualizations suggest that rate of reinforcement is the primary variable controlling the strength or levels of induced behavior. The operant response requirement within the schedule context has not been extensively studied as a determinant of induced responding. In the present study, levels of induced attack by food-deprived pigeons against restrained conspecifics were compared during response-dependent and response-independent schedules of food presentation equated or yoked interval-by-interval for reinforcement frequency. Experiment 1 compared levels of attack induced by fixed-ratio schedules of key pecking and yoked "matched-time" schedules. Experiment 2 similarly compared chained fixed-ratio 1 fixed-ratio 74 and yoked chained matched-time matched-time schedules. In both experiments, the response-dependent schedules generated greater levels (amount and probability) of induced attack than the response-independent time-based schedules. Thus, the ratio response requirement may be an important determinant of levels of induced responding, and the lower levels of attack observed during the response-independent condition may not be due to the absence of stimuli predicting food presentations. It is concluded that rate of reinforcement is not the sole variable determining levels of induced responding and that response-based and time-based schedules differ in their generation of induced responding.     

Choice between sequences of fixed-ratio schedules: effects of ratio values and probability of food delivery.

Pigeons were exposed to schedules of food delivery that consisted of two sequential fixed ratios. When alternative sequences provided two food deliveries per 50 responses, the schedule with the shorter initial fixed-ratio value was consistently preferred. Progressively reducing from 1.0 to .25 the probability of food delivery following completion of the second fixed ratio of the sequence with the shorter initial fixed ratio did not reduce preference for this sequence. Moreover, the sequence with the shorter initial fixed ratio also was preferred when the probability of food delivery following completion of the initial ratio in that sequence was progressively reduced from 1.0 to .5, although preference shifted to the alternative when the probability was reduced to 0. These findings suggest that the length of the initial fixed ratio was a primary determinant of choice. Subsequent manipulations demonstrated, however, that when the initial fixed ratios of the two alternatives were equal, changes in the ratio value and probability of food delivery following completion of the second fixed ratio lawfully affected choice.     

Reinforcing the absence of fixed-ratio performance

Pigeons received food for key pecking according to a fixed-ratio schedule, while, at the same time, food also was available for not pecking for a specified time. With a fixed ratio of 60, responding was not affected by not-pecking times of 80 or 40 seconds, and was eliminated completely at 10 seconds. With ratios of 180, pecking stopped with not-pecking times of 80 seconds or less; with ratios of 300, it stopped at 120 seconds or less. Not-responding schedules produced steady-state performance immediately following contact with the schedule. With return to the fixed-ratio schedule alone, response rate sometimes was elevated temporarily. When response-independent food presentations replaced the not-pecking schedule, response rate often was enhanced, and the ratio pattern was lost. Only the highest densities of food delivery eliminated responding, even with a fixed ratio of 300. In general, the effects corresponded to those of punishment, except that contrast had appeared both during and after punishment, and now appeared only after the response elimination procedure was suspended.     

Conjunctive schedules of reinforcement: III. A fixed-interval adjusting fixed-ratio schedule

Key pecking of three pigeons was studied under a conjunctive schedule that specified both a fixed-interval and an adjusting fixed-ratio requirement. The fixed-interval schedule was 6 min for one pigeon and 3 min for the other two. The size of the ratio requirement was determined within each cycle of the fixed interval by the duration of the pause before responding began. The fixed-ratio value was at maximum at the start of each fixed interval and decreased linearly until the first response occurred (adjusting fixed-ratio schedule). A peck produced food when the number of responses remaining on the fixed-ratio schedule was completed and when the fixed interval had elapsed. If no response occurred during the interval, the fixed-ratio requirement decreased to one and a single response after the interval elapsed produced food. The initial value of the adjusting fixed-ratio schedule was studied over a range of 0 to 900. Increases in the adjusting fixed-ratio schedule to about 300 responses increased both pause duration and running response rate and also modified the pattern of responding from that obtained under the fixed-interval schedule. Higher values of the adjusting fixed ratio generally decreased pause duration and running response rate and also disrupted responding. Interreinforcement time under the conjunctive schedule was increased substantially when the adjusting fixed-ratio size exceeded 300 responses.     

The effects of chlorpromazine and imipramine on rate and stimulus control of matching to sample.

Pigeons were trained to perform simultaneous, two-color matching to sample under a multiple fixed-ratio fixed-interval schedule of food presentation. The sequence terminating with a peck on the matching key (a "match") was treated as a unit, analogous to a single key peck in conventional schedules. Except for intermittent reinforcement of matches, no consequent stimulus distinguished matches from mismatches (sequences terminating with pecks on the nonmatching key). The pattern of matches during nondrug sessions resembled that of simpler operants maintained by similar schedules. Matches increased in rate toward the end of both components; mismatch rates increased more slowly. Imipramine increased the rate of mismatches, disrupted schedule patterning, and lowered accuracy in a dose-dependent fashion. Chlorpromazine lowered the overall rate of matches but affected schedule patterns and accuracy less than imipramine. The types of errors during drug sessions were not systematically related to the types of errors that appeared during nondrug sessions. Stimulus control was evaluated for each of the four possible color configurations and was found to be by the entire configuration of colors, not simply by the color of the sample.     

Choice between fixed-interval schedules: Graded versus step-like choice functions

Pigeons chose between two fixed-interval schedules of food reinforcement. A single peck on one of two lighted keys started the fixed-interval schedule correlated with that key. The schedule had to be completed before the next choice opportunity. The durations of the fixed intervals were varied over conditions from 15 s to 40 s. To maximize the rate of reinforcement, the pigeons had to choose exclusively the shorter of the two schedules. Nevertheless, choice was not all-or-none. Instead, relative choice, and the rates of producing the fixed intervals, varied in a graded fashion with the disparity between the two schedules. Choice ratios under this procedure (single response to choose) were highly sensitive to the ratios of the fixed-interval schedules.     

Conditioned reinforcement versus time to reinforcement in chain schedules.

Pigeons were trained on three-component chain schedules in which the initial component was either a fixed-interval or variable-interval schedule. The middle and terminal components were varied among fixed-interval fixed-interval, variable-interval variable-interval, and an interdependent variable-interval variable-interval schedule in which the sum of the durations of the two variable-interval components was always equal to the sum of the fixed-interval fixed-interval components. At issue was whether the response rate in the initial component was controlled by its time to primary reinforcement or by the temporal parameters of the stimulus correlated with the middle terminal link. The fixed-interval initial-link schedule maintained much lower response rates than the variable-interval initial-link schedule regardless of the schedules in the middle and terminal links. Nevertheless, the intervening schedules played some role: With fixed-interval schedules in the initial links, response rates were consistently highest with independent variable-interval schedules in the middle and terminal links and intermediate with the interdependent variable-interval schedules; these initial-link differences were predicted by the response rates in the middle link of the chain. With variable-interval schedules in the initial links, response rates were lowest with the fixed-interval fixed-interval schedules following the initial link and were not systematically different for the two types of variable-interval variable-interval schedules. The results suggest that time to reinforcement itself accounts for little if any variance in initial-link responding.