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.     

The effects of punishment intensity on squirrel monkeys

Responses of squirrel monkeys were maintained by a variable-interval schedule of food reinforcement. Concurrently, punishment consisting of a brief electric shock followed each response. As has been found for pigeons and rats, punishment did not produce extreme, all-or-none reactions. By gradually increasing the punishment intensity it was possible to produce response rates intermediate to no suppression and complete suppression. Similarly, the moment-to-moment response rate was free of extreme fluctuations. A “warm-up” effect occurred in which the punished responses were especially suppressed during the initial part of a session. The pre-punishment performance was negatively accelerated within a session, and punishment reduced the degree of negative acceleration. When punishment was discontinued, responding recovered immediately except when suppression had been complete or prolonged. When the punishment intensity was decreased gradually, more suppression resulted at a given intensity than when intensity was increased gradually. This suggests a “behavioral inertia” effect wherein behavior at a new punishment intensity is biased toward the behavior at the previous value. A corollary generalization is that the larger the change in intensity, the less the behavior at the new value will be biased toward the behavior at the previous value.     

Choice for signalled over unsignalled shock as a function of shock intensity

Choice between a signalled shock schedule and an unsignalled one was examined at various shock intensities. Three rats were given the opportunity to change from the unsignalled schedule to the signalled one at intensity values between 0.15 mA and 1.0 mA. Steps were usually 0.15 mA and both ascending and descending series were given. For two other rats, shock intensity increased from 0.20 mA to 1.0 mA in 0.20-mA increments; for two additional rats, shock intensity was first 3.0 mA and was then reduced to 1.0 mA. Subjects tended to remain in the unsignalled schedule at the lower shock intensities, but spent most of each session under the signalled schedule at the higher intensities (1.0 mA and 3.0 mA). In addition, the time spent in the signalled schedule tended to vary systematically with shock intensity over at least part of the range of intensity values. It was concluded that the relationship between shock intensity and choice behavior is similar to the relationship between intensity and behavior in procedures involving avoidance, escape, and punishment.     

The influence of positive and negative reinforcement on selective attention in the rat

In Experiments 1 and 2 rats were trained under two multiple schedules of reinforcement. In one, bar pressing during a tone-light compound stimulus was reinforced under a variable-interval food reinforcement schedule. In the other multiple schedule, bar pressing avoided grid shock on a free-operant schedule. In both multiple schedules, a discrimination was maintained by an extinction schedule that was operative during the absence of the tone-light compound. In Experiments 1 and 2 the intensity of the tone-light compound was manipulated over three levels. Subsequent extinction tests revealed that light was attended to, almost exclusively of the tone, when food reinforcement had maintained bar pressing. On the other hand, the tone gained considerable attentional control under the shock avoidance schedule. This stimulus-reinforcer interaction was maintained for all three levels of the compound intensity. In Experiment 3 it was investigated whether this interaction was associative by presenting shock during the absence of the tone-light compound when food reinforcement maintained responding, and food during the absence of the compound when shock avoidance maintained responding. Since both food and shock were presented during a single session for both schedules, nonassociative effects of the reinforcing stimuli were equivalent across the schedules. Nevertheless, the stimulus-reinforcer interaction was maintained, indicating that the interaction was an associative effect.     

Conditioned suppression of free-operant avoidance

The responses of white rats were maintained on an unsignalled free-operant avoidance schedule. Superimposed on the avoidance schedule was a blinking white light followed immediately by response-independent electric shock. Duration of the light stimulus was either 1 or 3 min. Avoidable shock was 1.5 mA; response-independent shock was 7.5 mA. Suppression of responding during the light stimulus (both durations) developed over sessions. Responding immediately following the response-independent light-shock sequence was neither suppressed nor accelerated. The similarity is noted between the present result and findings of “positive conditioned suppression”.     

Partial avoidance contingencies: Absolute omission and punishment probabilities

Avoidance contingencies were defined by the absolute probability of the conjunction of responding or not responding with shock or no shock. The “omission” probability (ρ₀₀) is the probability of no response and no shock. The “punishment” probability (ρ₁₁) is the probability of both a response and a shock. The traditional avoidance contingency never omits shock on nonresponse trials (ρ₀₀=0) and never presents shock on response trials (ρ₁₁=0). Rats were trained on a discrete-trial paradigm with no intertrial interval. The first lever response changed an auditory stimulus for the remainder of the trial. Shocks were delivered only at the end of each trial cycle. After initial training under the traditional avoidance contingency, one group of rats experienced changes in omission probability (ρ₀₀>0), holding punishment probability at zero. The second group of rats were studied under different punishment probability values (ρ₁₁>0), holding omission probability at zero. Data from subjects in the omission group looked similar, showing graded decrements in responding with increasing probability of omission. These subjects approximately “matched” their nonresponse frequencies to the programmed probability of shock omission on nonresponse trials, producing a very low and approximately constant conditional probability of shock given no response. Subjects in the punishment group showed different sensitivity to increasing absolute punishment probability. Some subjects decreased responding to low values as punishment probability increased, while others continued to respond at substantial levels even when shock was inevitable on all trials (noncontingent shock schedule). These results confirm an asymmetry between two dimensions of partial avoidance contingencies. When the consequences of not responding included occasional omission of shock, all subjects showed graded sensitivity to changes in omission frequency. When the consequences of responding included occasional shock delivery, some subjects showed graded sensitivity to punishment frequency while others showed control by overall shock frequency as well.     

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.     

Persistent behavior maintained by unavoidable shocks

Squirrel monkeys were trained on a multiple schedule in which 10-min periods on a continuous shock avoidance schedule, indicated by a yellow light, alternated with 10-min periods on a 1.5-min variable interval schedule of food reinforcement (VI 1.5). A white light indicated that VI 1.5 was in effect, except for the middle 2 min of the period on VI 1.5, in which a blue light appeared and terminated with the delivery of a 0.5-sec unavoidable shock. Stable response rates developed in the avoidance and VI 1.5 components. However, the highest response rates occurred in the blue, preshock stimulus. A series of experiments showed that responding in the blue stimulus persisted even when responding had been extinguished on both the VI schedule of food reinforcement and the shock avoidance schedule. Responding in the blue stimulus ceased when the blue stimulus terminated without shock or when it terminated with a response-contingent shock. Each time responding ceased, it was restored by terminating the blue stimulus with an unavoidable shock. When the blue stimulus was on throughout each session and unavoidable shocks were delivered at regular 10-min intervals, responding was well maintained. These results show that in monkeys that have been trained on a continuous avoidance schedule, unavoidable shocks can maintain responding even under conditions where responses have no programmed consequences.     

Development of a complex multiple schedule in the chimpanzee

The development of chimpanzee behavior on a four-component, three-lever multiple schedule is described. Component schedules included the Sidman avoidance procedure with a concurrent discriminated avoidance schedule on a second lever, fixed ratio performance for food, differential reinforcement of low rate for water requiring a dual response chain, and a symbol discrimination task for continuous food reinforcement using three levers. The avoidance component of this schedule was employed during the January 31, 1961 suborbital space flight of the chimpanzee “Ham.” On November 29, 1961, the chimpanzee “Enos” performed on the multiple schedule during three orbits around the earth in a Mercury capsule.     

Effects of chlorpromazine on appetitive and aversive components of a multiple schedule

A multiple schedule having both an appetitive and an avoidance component was maintained in two dogs to create a complex behavioral base line for observing the effects of chlorpromazine. Both soluble and “Spansule” chlorpromazine generated similar functions relating drug dose to measures of behavioral output. Although the dose ranges and the drugging procedures differed markedly for the different preparations of CPZ, the functions generated were comparable. There was no evidence that chlorpromazine had a differential depressing effect as a function of type of reinforcement. At low doses, rates of responding on the food-reinforced components increased slightly, whereas rates on the avoidance components remained relatively unchanged. At higher doses, both components showed an approximately equal depression of responding. These results are discussed with reference to some of the logical and experimental difficulties inherent in making comparisons across components of a multiple schedule and across schedules in general.     

Continuous punishment of free-operant avoidance in the rat

Three groups of albino rats were trained under a free-operant avoidance (Sidman) procedure with equal shock-shock and response-shock intervals. After stable performance was achieved, the animals were concurrently exposed to a brief electric shock after each response. The procedures were as follows: Punishment Schedule I: punishment shock was introduced at an intensity approximately one quarter that of avoidance shock; increments of nearly this same size were made as stable performance was achieved at succeeding punishment shock intensities. Punishment Schedule II: punishment shock was introduced at approximately one-half the intensity of avoidance shock; after stable performance, punishment shock was increased to the same intensity as avoidance shock. Punishment Schedule III: punishment shock was introduced and maintained at the same intensity as avoidance shock. Punishment was continued for all groups until one of two suppression criteria was attained. All animals made fewer responses and received more avoidance shocks as a function of increasing punishment shock. Half of the animals under Punishment Schedule I required punishment shock higher than avoidance shock to meet their assigned suppression criterion. A comparison of all procedures showed that suppression was greater when punishment shock was initially at high intensity.     

Competitive fixed-interval performance in humans

Two persons responded in the same session in separate cubicles, but under a single schedule of reinforcement. Each time reinforcement was programmed, only the first response to occur, that is, the response of only one of the subjects, was reinforced. “Competitive” behavior that developed under these conditions was examined in three experiments. In Experiment 1 subjects responded under fixed-interval (FI) 30-s, 60-s, and 90-s schedules of reinforcement. Under the competition condition, relative to baseline conditions, the response rates were higher and the pattern was “break-and-run.” In Experiment 2, subjects were exposed first to a conventional FI schedule and then to an FI competition schedule. Next, they were trained to respond under either a differential-reinforcement-of-low-rate (DRL) or fixed-ratio (FR) schedule, and finally, the initial FI competition condition was reinstated. In this second exposure to the FI competition procedure, DRL subjects responded at lower rates than were emitted during the initial exposure to that condition and FR subjects responded at higher rates. For all subjects, however, responding gradually returned to the break-and-run pattern that had occurred during the first FI competition condition. Experiment 3 assessed potential variables contributing to the effects of the competitive FI contingencies during Experiments 1 and 2. Subjects were exposed to FI schedules where (a) probability of reinforcement at completion of each fixed interval was varied, or (b) a limited hold was in effect for reinforcement. Only under the limited hold was responding similar to that observed in previous experiments.     

The effects of cocaine on behavior maintained by timeout from avoidance.

Rats were trained on concurrent schedules under which responses on one lever postponed shock (avoidance) and responses on the other lever produced brief (2-min) periods of signaled timeout from avoidance. For 6 rats, timeout from avoidance was programmed on a variable-interval 45-s schedule that generally resulted in rates that were lower than those on the avoidance lever. For another 6 rats, timeout was arranged on a variable-ratio 15 schedule that produced higher baseline rates. Cocaine (3 to 40 mg/kg) produced large, dose-dependent increases in behavior maintained by timeout in both groups of rats. Avoidance responding was also generally increased by cocaine, but the increases were of lesser magnitude. Increases in response rates were seen across a broad range of doses on behavior maintained by either interval or ratio schedules, an outcome that was unexpected on the basis of most studies of cocaine on food-maintained behavior. These results were similar to those of previous studies of the effects of amphetamine on behavior maintained by timeout from avoidance and suggest that stimulant drugs affect behavior maintained under a shock-postponement schedule differently than they affect behavior maintained by timeout from avoidance.     

Effects of response-shock interval and shock intensity on free-operant avoidance responding in the pigeon

Two experiments investigated free-operant avoidance responding with pigeons using a treadle-pressing response. In Experiment I, pigeons were initially trained on a free-operant avoidance schedule with a response-shock interval of 32 sec and a shock-shock interval of 10 sec, and were subsequently exposed to 10 values of the response-shock parameter ranging from 2.5 to 150 sec. The functions relating response rate to response-shock interval were similar to the ones reported by Sidman in his 1953 studies employing rats, and were independent of the order of presentation of the response-shock values. Shock rates decreased as response-shock duration increased. In Experiment II, a free-operant avoidance schedule with a response-shock interval of 20 sec and a shock-shock interval of 5 sec was used, and shock intensities were varied over five values ranging from 2 to 32 mA. Response rates increased markedly as shock intensity increased from 2 to 8 mA, but rates changed little with further increases in shock intensity. Shock rates decreased as intensity increased from 2 to 8 mA, and showed little change as intensity increased from 8 to 32 mA.     

Maintenance and suppression of responding under schedules of electric shock presentation

In squirrel monkeys previously trained under a continuous avoidance schedule, characteristic patterns of responding were maintained under a 3-min variable-interval schedule of shock presentation (response-produced shock). Responding in the presence of a periodically presented stimulus, the termination of which coincided with the delivery of a response-independent electric shock (Estes-Skinner procedure), was not reliably affected. When shocks followed every response during certain signalled portions of the session, and were presented under the variable-interval schedule during the rest of the session (multiple 1-response fixed-ratio, 3-min variable-interval schedule of shock presentation), responding was suppressed during the fixed-ratio component and maintained during the variable-interval component. Environmental consequences do not have immutable properties, and may either support or suppress behavior, depending on the schedule of presentation.     

Running-wheel activity and avoidance in the mongolian gerbil

In the first of two experiments, running-wheel activity and unsignaled (Sidman) avoidance were studied in gerbils and albino rats. All gerbils ran at higher rates than any of the four rats studied. Under the avoidance procedure, four rats developed effective responding; the other two performed much less successfully. While avoidance developed more slowly in the gerbils, all showed asymptotic performance as effective as the four superior rats. The rats showed a consistent warm-up effect, receiving 60% to 80% of the total shocks in the initial third of the session. The gerbils displayed no warm-up, with shocks being evenly distributed over the session. Warm-up in the rats was not related to either response rate or to how effectively the animal was avoiding. When shock was removed, extinction occurred more rapidly in the rats than the gerbils. In the second experiment, which involved two-way shuttle avoidance, gerbils and albino mice quickly acquired the response. All animals met a criterion of 90% avoidance over 80 trials.     

RESPONDING MAINTAINED UNDER INTERMITTENT SCHEDULES OF ELECTRIC-SHOCK PRESENTATION: “SAFETY” OR SCHEDULE EFFCTS?

Four experiments were conducted in which lever pressing by squirrel monkeys was maintained under multiple, mixed, or chained schedules of electric-shock presentation. In the first two experiments, a multiple schedule was employed in which a fixed-interval schedule of shock presentation alternated with a signaled two-minute component. Initially, no events were scheduled during the two-minute component (a safety period). In the first experiment, the safety period was “degraded” by introducing and systematically increasing the frequency of periodic shocks presented during that component. In the second experiment, the proportion of overall safe time to unsafe time was decreased by decreasing the value of the fixed-interval schedule while holding constant shock frequency during the two-minute component. In the third experiment, the overall arrangement was changed from a multiple to a mixed schedule in an attempt to determine whether fixed-interval responding would be maintained when a single exteroceptive stimulus was associated with both components. In the fourth experiment, the overall arrangement was changed from a multiple to a chained schedule in an effort to determine whether fixed-interval responding would be maintained when its consequence was presentation of a signaled “unsafe” period. Fixed-interval responding was well maintained under all experimental conditions; the varied relationships obtained lend more support to conceptualizations of shock-maintained behavior as exemplifying schedule-controlled behavior than to suggestions that such behavior may be readily accounted for by “safety theory.”     

Length of experimental session as a parameter of response rate under a non-discriminated

Following the stabilization of response rate under an avoidance schedule which was defined by two temporal parameters, the shock-shock interval and the interval by which each response postponed the onset of shock, the length of the experimental session was changed. It was found that after the subjects had been exposed to a longer session of avoidance schedule, their rates of response were considerably increased without a corresponding reduction in the number of shocks received.

In recent years considerable use has been made of an avoidance training technique in which the performance of the response functions to postpone the onset of an aversive stimulus, usually shock, by a fixed period. In the absence of the required response the aversive stimulus is programmed to occur at regular intervals. Experiments by Sidman (1953) have shown that the critical independent variable controlling the rate of avoidance response, is the shock postponement interval (R*S). All other things being equal, the rat in the lever pressing situation will respond at a rate which is inversely related to the R*S interval, low intervals generating high response rates and high intervals generating low response rates. However, under very low values of R*S, the response rate may break down altogether. The animal then receives shock at the rate determined by the shock-shock interval parameter.

As a result of an apparatus failure, Sidman, Herrnstein and Conrad (1957) discovered that the response rate can also be increased by occasionally shocking the animal in spite of its avoidance responses. An apparatus failure has also been responsible for the isolation of yet another parameter of response rate in the shock-postponement avoidance situation and is reported here. Briefly, it was found that a change in the duration of an experimental session influences the response rate on subsequent sessions.     

Punishment: the interactive effects of delay and intensity of shock

A discrete-trial punishment procedure, with rats, was used to examine how delay-of-shock intervals of 0 to 28 sec and shock intensity interact to decrease the frequency and increase the latency of a positively reinforced response. For delay-of-shock intervals of 0, 7, 14, and 28 sec, there was a range of shock intensities, for some subjects, over which the punishing effect of shock was an increasing, monotonic function of shock intensity. For other subjects this transition was abrupt. Functions relating response frequency and latency measures to shock intensity were displaced toward higher values on the shock intensity axis with an increase in delay-of-shock interval. The effects of “gradual” and “abrupt” introduction to “severe” shock, as well as re-exposure to previously used shock intensities, were examined under both the immediate and delay-of-shock conditions. With delay-of-shock intervals of 7, 14, or 28 sec, shock intensities of approximately 0.50 milliamperes or greater were necessary to decrease substantially the number and increase the latency of the lever-pressing response. For the immediate punishment group this intensity was approximately 0.20 ma. These facts were related to Annau and Kamin's (1961) conditioned emotional response experiment in which a shock intensity of 0.49 ma or greater was required to suppress the rate of a positively reinforced response.     

Immobility as an avoidance response, and its disruption by drugs

Much of the available literature on avoidance behavior is based on responses which require the animal to run, lever-press, or to make some active response to avoid noxious stimulation. The purpose of Experiment I reported in this paper was to determine whether animals can learn to sit or stand motionless in order to escape or avoid electric shock. Five experimental rats were given escape-avoidance training, while five yoked control animals received electric shocks without any response-related contingency. It was shown that an immobility avoidance response, as distinct from the unconditioned “freezing” response to shock, can be trained. The results of Experiment II (30 rats) revealed that this response is more readily acquired at higher shock intensities than at lower ones, provided escape by jumping is prevented at the high shock intensities. The effects of six doses of each of three drugs on the immobility avoidance response were studied in Experiment III (13 rats). Methylphenidate, chlorpromazine, and imipramine all produced a decrement in the immobility response, but the pattern and amount of the effects of the three drugs were quite different, one from the other. The implications of these findings for a general theory of avoidance behavior and for drug screening are discussed.