Effects of diazepam on schedule-controlled and schedule-induced behavior under signaled and unsignaled shock.

Schedule-controlled lever pressing and schedule-induced licking were studied in rats under a multiple fixed-interval fixed-interval schedule of food reinforcement upon which was superimposed a multiple variable-time variable-time schedule of electric-shock delivery. Shocks were signaled in one component of the multiple schedule and unsignaled in the other. The effects of diazepam upon the suppression of behavior during the signal (conditioned suppression) and during signaled and unsignaled shock (differential suppression) were studied under several shock intensities (Experiment 1) and at increased body weight (Experiment 2). In each study, diazepam led to dose-dependent increases in the rate of pressing and licking during signaled and unsignaled shock, but had little effect on conditioned suppression. the rate-enhancing effects of diazepam depended upon the intensity of shock, nature of the response, and whether or not shocks were signaled. The data was discussed in terms of (1) implications for understanding the effects of signaled and unsignaled shock on behavior, (2) the effects of diazepam on behavior suppressed by response-independent shock, and (3) comparison between operant and schedule-induced behavior.     

Effects of response-dependent and independent electric shock on schedule-induced polydipsia

In Experiment I, lever pressing by rats was maintained by the delivery of food pellets under a 45-sec fixed-interval schedule. Fixed-time 180-sec and fixed-interval 180-sec schedules of shock delivery were systematically superimposed on the baseline food schedule to study effects on schedule-induced water intake. Response-dependent shock had little, if any, effect on water intake, whereas shocks independent of lever pressing attenuated fluid intake. In Experiment 2, rats received food pellets under a fixed-time 60-sec schedule. Electric shock delivered concurrently under a variable-time 180-sec schedule, but never while the animal was licking or within 5 sec after licking terminated, led to similar attenuation of water intake. These findings suggest that schedule-induced polydipsia is sensitive to differences in the functional properties of response-independent and dependent electric shock.     

Effects of signaled and unsignaled shock on schedule-controlled lever pressing and schedule-induced licking: Shock intensity and body weight

Schedule-controlled lever pressing and schedule-induced licking were studied in rats under a multiple fixed-interval fixed-interval schedule of food reinforcement. Following acquisition of stable rates of pressing and licking, a multiple variable-time variable-time schedule of electric-shock delivery was superimposed upon the baseline schedule. In only one component of the multiple schedule, a 5-sec stimulus preceded each shock (signaled shock). In the other component shock was unsignaled. Several shock intensities (Experiment 1) and body weights (Experiment 2) were studied. Lever pressing and licking were affected similarly by experimental manipulations, although with parametric differences. Depending upon shock intensity and body weight, rates of lever pressing and licking were hardly suppressed, suppressed primarily in the unsignaled shock component (differential suppression), or markedly suppressed in both components. Differential suppression during components with signaled and unsignaled shock and conditioned suppression of responding during the preshock stimulus appeared not to be functionally related. Differential suppression depended more on the discriminability of shock-free time, and on shock intensity, body weight, and the type of response than on the “preparatory” behavior preceding shock.     

Intercurrent and reinforced behavior under multiple spaced-responding schedules

Lever pressing in rats was reinforced with food under a multiple spaced-responding schedule. A lever, food cup, and drinking tube were mounted in a running wheel so that lever pressing, running, and licking could be recorded. Running and licking had no scheduled consequences. Lever pressing was reinforced under a multiple schedule with three spaced-responding components and an extinction component. Each component was associated with a different auditory stimulus. Spaced-responding components reinforced only lever presses terminating interresponse times equal to or greater than 10, 20, or 60 sec, respectively. Rates of lever pressing, reinforcement, and licking all decreased as schedule parameter increased. Efficiency of spaced responding, as measured by reinforcements per response, also decreased. Rate of wheel running either increased or increased and then decreased with increasing schedule parameter. Individual running rates differed substantially. Neither licking nor running rate correlated with individual differences in efficiency. Analysis of conditional probabilities among the several response classes showed that, as the schedule requirement increased, the probability of running after a lever press increased and the probability of licking after a lever press decreased. After reinforcement, one subject always pressed the lever next. In the other subjects, the conditional probability of lever pressing, given reinforcement, increased while the probability of licking, given reinforcement, decreased with increasing schedule requirement. Results are discussed in relation to the concepts of schedule-induced and mediating behavior.     

Effects of fixed-time shocks and brief stimuli on food-maintained behavior of rats

When a fixed-time schedule of shocks was presented to rats lever pressing for food on a random-interval schedule, a pattern of behavior developed with a high rate of pressing after shock declining to near zero before the next shock was delivered. Once this pattern had stabilized, one-quarter of the shocks were replaced with brief auditory stimuli (tones) in a random sequence. Tone maintained behavior similar to shock, although tone was never paired with shock. Both tone and shocks elicited responding when presented at various times as probe stimuli, and responding was usually totally suppressed if neither stimulus occurred at the beginning of the fixed-time interval. When other stimuli were paired with tone and shock, only those paired with tone gained discriminative control and elicited responding. These findings suggest that stimuli that signal a shock-free, or safe, period will maintain the pattern of behavior generated by shock on a fixed-time schedule. There is a parallel between this phenomenon and the control of behavior on second-order schedules of positive reinforcement with nonpaired brief stimuli.     

Schedule-induced licking during multiple schedules

Schedule-induced polydipsia was studied in rats bar pressing under two-component multiple schedules of food reinforcement. The first component of the multiple schedule was a variable-interval 1-min schedule throughout the experiment. The schedule comprising the second component was varied over blocks of sessions in terms of rate and magnitude of reinforcement, and was either variable-interval 3-min (one pellet), variable-interval 3-min (three pellets), variable-interval 1-min (one pellet), or extinction. Water intake per session varied with the rate of reinforcement in the schedule comprising the second component and was highest when the schedule was variable-interval 1-min. Both bar-pressing behavior and licking behavior showed behavioral interactions between the two components of the multiple schedules. With magnitude of reinforcement held constant, a matching relationship was observed between lick rate and reinforcement rate; the relative frequency of licks in the constant component matched the relative frequency of reinforcement in that component. Bar pressing, however, showed only a moderate degree of relativity matching. During the schedule-induced licking, a burst of licking followed each delivery of a pellet (post-prandial drinking). The duration of these bursts of licking was observed to be a function of the inter-reinforcement interval.     

Schedule-induced defecation by rats during ratio and interval schedules of food reinforcement.

Lever pressing in rats was maintained by continuous and intermittent schedules of food while defecation was monitored. In Experiment 1, reinforcement densities were matched across variable-ratio and variable-interval schedules for three pairs of rats. Defecation occurred in all 3 rats on the variable-ratio schedule and in all 3 rats on the yoked variable-interval schedule. In Experiment 2, fixed-ratio and fixed-interval schedules with similar reinforcement densities maintained lever pressing. Defecation occurred in 3 of 4 rats on the fixed-ratio schedule and in 4 of 4 rats on the fixed-interval schedule. Almost no defecation occurred during continuous reinforcement in either experiment. These results demonstrate that defecation may occur during both ratio and interval schedules and that the inter-reinforcement interval is more important than the behavioral requirements of the schedule in generating schedule-induced defecation.     

A direct comparison of four methods for eliminating a response

Thirty-two rats pressed one lever (lever A) on a VI 30-sec schedule of food reinforcement and were then shifted to one of four procedures for eliminating the lever A response: extinction, differential reinforcement of other behavior, reinforcement of a different response (pole pushing), and reinforcement of a similar response (pressing lever B). Effectiveness of a response-elimination procedure was measured by (1) how quickly lever A response rate fell to a low level when the procedure was in effect, (2) how much lever A responding recovered when the procedure was discontinued, and (3) how resistant lever A responding was to reinstatement when the VI reinforcement schedule was reimposed. No one method was superior by all three measures. Extinction produced the most variable behavior, while differential reinforcement of other behavior produced the least. Reinforcing alternative behavior produced the greatest recovery in the original lever A response when the response-elimination procedure was discontinued.     

Effects of d-amphetamine and caffeine on schedule-controlled and schedule-induced responding.

The effects of d-amphetamine and caffeine were studied on rates and patterns of lever pressing and schedule-induced licking under fixed-interval schedules of food pellet presentation. In addition, the effects of caffeine were studied on lever pressing and licking under a multiple fixed-ratio fixed-interval schedule. Caffeine reduced mean overall rates of licking at lower doses than it reduced mean overall rates of pressing under the fixed-interval schedules, but the effects of caffeine on both licking and lever pressing depended largely on the control rate of responding. d-Amphetamine reduced mean overall rates of lever pressing and licking at about the same dose, but the effects of d-amphetamine also were a function of the control rate of responding.     

Conditioned suppression and conditioned enhancement with the same positive UCS: an effect of CS duration

Previous experiments have shown that positively reinforced operant responding is suppressed during a conditioned stimulus terminated with an electric shock (conditioned suppression). In the present experiment, the conditioned stimulus was terminated with a positive unconditioned stimulus, and it was found that the duration of the conditioned stimulus was a key factor in determining whether response suppression or response enhancement was observed during the stimulus. The lever-pressing responses of rats were maintained by a variable-interval schedule of food reinforcement. While the rats were pressing the lever, a light was occasionally turned on, its offset coincident with a brief period of access to a sucrose solution. In consecutive blocks of sessions, the light duration was 40 sec, 12 sec, or 120 sec. Results showed that the rate of lever pressing was substantially suppressed during the 12-sec stimulus, slightly suppressed during the 40-sec stimulus, and enhanced during the 120-sec stimulus.     

Food-deprivation effects on punished schedule-induced drinking in rats.

Food-deprived rats (at 80% of their free-feeding weights) were exposed to a fixed-time 60-s schedule of food-pellet presentation and developed schedule-induced drinking. Lick-dependent signaled delays (10 s) to food presentation led to decreased drinking, which recovered when the signaled delays were discontinued. A major effect of this punishment contingency was to increase the proportion of interpellet intervals without any licks. The drinking of yoked control rats, which received food at the same times as those exposed to the signaled delay contingency (masters), was not consistently reduced. When food-deprivation level was changed to 90%, all master and yoked control rats showed decreases in punished or unpunished schedule-induced drinking. When the body weights were reduced to 70%, most master rats increased punished behavior to levels similar to those of unpunished drinking. This effect was not observed for yoked controls. Therefore, body-weight loss increased the resistance of schedule-induced drinking to reductions by punishment. Food-deprivation effects on punished schedule-induced drinking are similar to their effects on food-maintained lever pressing. This dependency of punishment on food-deprivation level supports the view that schedule-induced drinking can be modified by the same variables that affect operant behavior in general.     

A comparison of operant licking and lever pressing in the rat

Two groups of rats were trained on a signaled, free-operant, avoidance procedure to lick or to lever press in order to avoid shock while water-deprived or satiated and, in the case of licking, while ingesting deionized water, isotonic saline, or 10% sucrose. The most effective avoidance licking occurred while the rats were water-deprived and ingesting 10% sucrose. Water deprivation level had no effect on lever pressing for shock avoidance. Two other groups of rats were operantly conditioned to lick or to lever press for food pellets while waterdeprived or satiated and, in the case of licking, while ingesting deionized water or 10% sucrose. The most effective licking for food reinforcement occurred while the rats were water-deprived and ingesting 10% sucrose. Water deprivation level had no effect on lever pressing for food reinforcement. The data indicated that effective operant licking must be supported by factors related to water regulation and taste palatability.     

Schedules of food postponement: II. Maintenance of behavior by food postponement and effects of the schedule parameter

In Experiment I, food-deprived, feeder-trained squirrel monkeys pressed a lever to postpone brief electric shocks (Response-Shock=Shock-Shock interval=30 seconds). Forty-one three-hour sessions of shock postponement were followed by 120 sessions of concurrent shock and food postponement. The shock schedule was unchanged and the food schedule was Response-food interval–20 seconds, Food-food interval 10 seconds. After concurrent shock and food postponement, the shock schedule was discontinued and 40 sessions of food postponement ensued, followed by 53 sessions of extinction. After extinction, food postponement was resumed for 11 sessions. Stable responding with low food rates was maintained under food-postponement after the concurrent schedule. Responding decreased to low levels under extinction and recovered immediately to previous levels when the food-postponement schedule was re-instated. In Experiment II, a parameter of the food-postponement schedule was studied sequentially. Using the same subjects, the Response-food–Food-food interval was manipulated from four seconds to 80 seconds with several orders of presentation. Relations of response rates and food rates to the parameter were similar to those seen under shock postponement. Exposure to very short postponement times (four seconds), resulting in very high food rates, decreased but did not abolish subsequent responding at longer postponement times. Results are discussed from the point of view that reinforcing functions of stimuli consequent on responding depend on a prior history of scheduled contact with those stimuli.     

Responding under schedules combining response-dependent and response-independent shock delivery

Lever pressing of three squirrel monkeys with experience under continuous avoidance schedules was maintained by response-produced shock under a 5-minute variable-interval schedule. Responding decreased when half of the scheduled shocks were delivered independently of lever pressing and decreased further when all shocks were independent of lever pressing. Responding was lowest when all shocks were eliminated. When the proportion of response-dependent shocks increased, responding increased. This relation occurred even though the frequency and temporal distribution of shock delivery remained the same. Responding of two monkeys increased in a graded fashion as the frequency of shock was increased by arranging variable-time 5-minute, 2-minute, and 1-minute schedules jointly with the variable-interval 5-minute schedule. Thus, increasing the proportion of response-independent shocks decreased responding when the overall frequency of shocks stayed the same, but increased responding when the overall frequency of shock delivery increased.     

Schedule-induced biting under fixed-interval schedules of food or electric-shock presentation

Squirrel monkeys pressed a lever under fixed-interval schedules of food or of electric-shock presentation. Both schedules induced repeated biting on a latex hose. Whether lever pressing was controlled by food or by electric shock, a pattern of decreasing hose biting and increasing lever pressing occurred within fixed-interval cycles. As the fixed-interval duration was increased from 6 to 600 sec, average rates of lever pressing decreased under both schedules. Average rates of hose biting first increased with increasing parameter value, reaching a maximum at values that varied from 60 to 337 sec in different monkeys, and then declined at higher values. d-Amphetamine at appropriate doses increased overall rates of lever pressing maintained by food or by shock, but either did not affect or decreased overall rates of hose biting. When no timeout period occurred between fixed-interval cycles, the monkeys bit most frequently immediately after food or electric shock was presented. When there was a timeout period, hose biting began shortly after the start of the fixed-interval cycles, with little or no hose biting immediately after food or electric shock was presented. Most hose biting appeared to be schedule-induced rather than food- or shock-induced.     

Behavior simultaneously maintained by both presentation and termination of noxious stimuli

Lever pressing by two squirrel monkeys was maintained under a 3-minute variable-interval schedule of response-produced electric-shock presentation. At the same time, responding on a second lever was maintained under a 3-minute fixed-interval schedule of termination of the shock-presentation schedule and shock-correlated stimuli. Under the termination schedule, the first response after a 3-minute period produced a 1-minute timeout, during which no events occurred and responding had no scheduled consequence. Relatively high and constant rates of responding were maintained on the lever where responding produced shock. Lower rates and positively accelerated patterns of responding occurred on the lever where responding terminated the shock schedule. Thus, responding was simultaneously maintained by presentation of an event and by termination of a stimulus associated with that event. Rates and patterns of responding on each lever were reversed when the schedules arranged on each lever were reversed on two occasions. When shock intensity was increased from 0 to 10 mA, responding maintained both by presentation of shock and by termination of the shock schedule increased, but responding maintained by shock presentation increased to a greater extent. Positive and negative reinforcement, usually regarded as separate behavioral processes involving different events, can coexist when behavior is controlled by different contingencies involving the same event.     

Concurrent fixed-ratio and avoidance responding in the squirrel monkey

Squirrel monkeys maintained concurrent performances appropriate to a fixed-ratio schedule of food reinforcement on one lever and an avoidance schedule on a second lever. The overall rate of responding maintained by either schedule was not systematically affected when the other schedule was discontinued and its lever removed.     

Extinction of responding maintained by timeout from avoidance

The resistance to extinction of lever pressing maintained by timeout from avoidance was examined. Rats were trained under a concurrent schedule in which responses on one lever postponed shock on a free-operant avoidance (Sidman) schedule (response-shock interval = 30 s) and responses on another lever produced 2 min of signaled timeout from avoidance on a variable-ratio 15 schedule. Following extended training (106 to 363 2-hr sessions), two experiments were conducted. In Experiment 1 two different methods of extinction were compared. In one session, all shocks were omitted, and there was some weakening of avoidance but little change in timeout responding. In another session, responding on the timeout lever was ineffective, and under these conditions timeout responding showed rapid extinction. The within-session patterns produced by extinction manipulations were different than the effects of drugs such as morphine, which also reduces timeout responding. In Experiment 2 shock was omitted for many consecutive sessions. Response rates on the avoidance lever declined relatively rapidly, with noticeable reductions within 5 to 10 sessions. Extinction of the timeout lever response was much slower than extinction of avoidance in all 4 rats, and 2 rats continued responding at baseline levels for more than 20 extinction sessions. These results show that lever pressing maintained by negative reinforcement can be highly resistant to extinction. The persistence of responding on the timeout lever after avoidance extinction is not readily explained by current theories.     

Variable-ratio schedules of timeout from avoidance: effects of d-amphetamine and morphine.

Rats were trained on concurrent schedules in which pressing one lever postponed shock and pressing the other lever produced periods of signaled timeout from avoidance on variable-ratio schedules. These procedures generated high rates of timeout-reinforced responding and provided a baseline for studying the effects of drugs on behavior maintained by different types of negative reinforcement (shock postponement vs. timeout). Morphine (2.5 to 10.0 mg/kg) reduced behavior maintained by timeout at doses that increased or had no effect on avoidance responding. In contrast, d-amphetamine (0.125 to 2.0 mg/kg) produced large increases in timeout responding at doses that had minimal effect on avoidance in rats trained on variable-interval and variable-ratio schedules. Thus, the event-dependent effects of morphine, observed in previous studies in which timeout responding was maintained at low rates by interval schedules, were replicated with high timeout rates maintained by variable-ratio schedules. The effects of d-amphetamine could also be described as "event dependent" because timeout responding was stimulated more than avoidance regardless of the maintenance schedule or baseline rate.     

Preference for less frequent shock under fixed-interval schedules of electric-shock presentation.

Lever pressing by 2 squirrel monkeys was maintained under fixed-interval 6-min and fixed-interval 2-min schedules of electric-shock presentation. Preference for these schedules was assessed during three experimental phases. In all phases, responses on one lever produced shock according to one or the other fixed-interval schedule, and responses on a second, changeover, lever switched between schedules. The opportunity to change over was presented during separate choice periods (during which the fixed-interval schedules did not operate) that followed the first through fourth shocks in each schedule. If no changeover occurred during those choice periods, a changeover automatically occurred following the fifth shock. In Phase I, durations of the choice periods were fixed. In Phase II, the choice periods equaled a proportion of their respective fixed interval. During Phase III (completed with 1 monkey) a response on the changeover lever during a given choice period reinstated the most recent fixed interval, and a failure to respond resulted in a changeover. During each of these phases, distinct preferences developed for the 6-min schedule. These results suggest that the maintenance of lever pressing by fixed-interval presentation of electric shock may not be an example of positive reinforcement, and that the response-maintaining characteristics of shock presentation may derive from other properties of the schedule.