ASSESSING THE ROLE OF EFFORT REDUCTION IN THE REINFORCING EFFICACY OF TIMEOUT FROM AVOIDANCE

Rats responded on concurrent schedules of shock‐postponement or deletion (avoidance) and timeout from avoidance. In Experiment 1, 3 rats' responses on one lever postponed shocks for 20 s and responses on a second lever produced a 1‐min timeout according to a variable‐interval 45‐s schedule. Across conditions, a warning signal (white noise) was presented 19.5 s, 16 s, 12 s, 8 s, or 4 s before an impending shock. Raising the duration of the warning signal increased both avoidance and timeout response rates. Timeout responding, although positively correlated with avoidance responding, was not correlated with the prevailing shock rate. In Experiment 2, 3 rats' responses on one lever deleted scheduled shocks according to a variable‐cycle 30‐s schedule and responses on a second lever produced a 2‐min timeout as described above. After this baseline condition, the avoidance lever was removed and noncontingent shocks were delivered at intervals yoked to the receipt of shocks in the baseline sessions. Timeout responding decreased when the avoidance lever was removed, even though the shock‐frequency reduction afforded by the timeout remained constant. These results suggest that a key factor in the reinforcing efficacy of timeout is suspension of the requirement to work to avoid shock, rather than the reduction in shock frequency associated with timeout.     

Reductions in shock frequency and response effort as factors in reinforcement by timeout from avoidance

Rats' presses on one lever canceled shocks programmed after variable cycles, while presses on a second lever occasionally produced a 2-min timout during which the shock-delection schedule was suspended and its correlated stimuli removed. These concurrent schedules of avoidance and timeout were embedded in a multiple schedule whose components differed, within and across conditions, in terms of the programmed shock rate associated with the shock-deletion schedule. Analyses based on the generalized matching law suggest that the reduction in the response requirement correlated with termination of the avoidance schedule was a more important factor in the reinforcing effectiveness of timeout than was shock-frequency reduction, at least in 2 of 3 rats. After training in each condition, responding on the timeout lever was extinguished by withholding timeouts in both components over seven sessions. Resistance to extinction varied directly with the rates of both shock-frequency reduction and avoidance-response reduction experienced during training. Although reduction in response effort appeared to dominate shock-frequency reduction in the maintenance of responding, neither factor had a clear advantage in predicting the course of extinction.     

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.     

Relative reinforcer magnitude under a nonindependent concurrent schedule of cocaine reinforcement in rhesus monkeys.

Lever pressing by three rhesus monkeys was maintained under a two-lever concurrent schedule of cocaine reinforcement. Responding on one lever (constant-dose lever) produced a constant dose of 0.05 or 0.1 mg/kg/injection arranged according to a variable-interval 1-min schedule. Responding on the other lever (variable-dose lever) produced a comparison dose of cocaine (0.013 to 0.8 mg/kg/injection), also under a variable-interval 1-min schedule. The two variable-interval schedules were made nonindependent by arranging that the assignment of a reinforcer by one schedule inactivated the second schedule until the assigned reinforcer had been obtained. This modification ensured that the two cocaine doses were obtained with approximately equal frequency, regardless of the distribution of the subject's responding. Preference, indicated by relative response frequency on the variable-dose lever, was almost always for the larger of the doses and was a monotonic function of the comparison dose, except at the highest doses. Preferences at the highest comparison doses may have resulted from the low overall response rates exhibited at these doses. Relative response frequencies on the variable-dose lever roughly matched relative reinforcer magnitude (mg/kg/injection available on the variable-dose lever divided by the sum of mg/kg/injections available on each lever).     

Lick-shock contingencies in the rat

Hungry rats were allowed to lick an 8% sucrose solution and then one of four lick-shock contingency conditions was superimposed on the licking baseline. These conditions were: free-operant avoidance, free shock, punishment, and no shock. From highest to lowest response rates, the groups fell in the order-avoidance, no shock, free shock, and punishment. Lick rates adjusted rapidly to introduction and removal of the contingencies. Post-shock responding was lowest in the punishment condition and highest in the free shock condition. No method was found simultaneously to equate shock frequency and separate response rates for the three shock contingency conditions. Only small, or no, reductions in shock rate occurred over sessions under the free-operant avoidance schedule when the shock-shock interval was 10 sec but large reductions occurred when the shock-shock interval was reduced to either 1 or 2 sec.     

SUPPRESSIVE AND FACILITATIVE EFFECTS OF SHOCK INTENSITY AND INTERRESPONSE TIMES FOLLOWED BY SHOCK

Although response‐dependent shock often suppresses responding, response facilitation can occur. In two experiments, we examined the suppressive and facilitative effects of shock by manipulating shock intensity and the interresponse times that produced shock. Rats' lever presses were reinforced on a variable‐interval 40‐s schedule of food presentation. Shock followed either long or short interresponse times. Shock intensity was raised from 0.05 mA to 0.4 mA or 0.8 mA. Overall, shock contingent on long interresponse times punished long interresponse times and increased response rates. Shock contingent on short interresponse times punished short interresponse times and decreased response rates. In Experiment 1, raising the range of interresponse times that produced shock enhanced these effects. In Experiment 2, the effects of shock intensity depended on the interresponse times that produced shock. When long interresponse times produced shock, low intensities increased response rates. High intensities decreased response rates. When short interresponse times produced shock, high shock intensities punished short interresponse times and decreased response rates more than low intensities. The results may explain why punishment procedures occasionally facilitate responding and establish parameters for future studies of punishment.     

HUMAN RESPONDING ON RANDOM‐INTERVAL SCHEDULES OF RESPONSE‐COST PUNISHMENT: THE ROLE OF REDUCED REINFORCEMENT DENSITY

An experiment with adult humans investigated the effects of response‐contingent money loss (response‐cost punishment) on monetary‐reinforced responding. A yoked‐control procedure was used to separate the effects on responding of the response‐cost contingency from the effects of reduced reinforcement density. Eight adults pressed buttons for money on a three‐component multiple reinforcement schedule. During baseline, responding in all components produced money gains according to a random‐interval 20‐s schedule. During punishment conditions, responding during the punishment component conjointly produced money losses according to a random‐interval schedule. The value of the response‐cost schedule was manipulated across conditions to systematically evaluate the effects on responding of response‐cost frequency. Participants were assigned to one of two yoked‐control conditions. For participants in the Yoked Punishment group, during punishment conditions money losses were delivered in the yoked component response independently at the same intervals that money losses were produced in the punishment component. For participants in the Yoked Reinforcement group, responding in the yoked component produced the same net earnings as produced in the punishment component. In 6 of 8 participants, contingent response cost selectively decreased response rates in the punishment component and the magnitude of the decrease was directly related to the punishment schedule value. Under punishment conditions, for participants in the Yoked Punishment group response rates in the yoked component also decreased, but the decrease was less than that observed in the punishment component, whereas for participants in the Yoked Reinforcement group response rates in the yoked component remained similar to rates in the no‐punishment component. These results provide further evidence that contingent response cost functions similarly to noxious punishers in that it appears to suppress responding apart from its effects on reinforcement density.     

Maintenance of responding by squirrel monkeys under a concurrent shock-postponement, fixed-interval shock-presentation schedule.

A chain-pulling response was initially developed under a shock-postponement (avoidance) schedule with two squirrel monkeys. Few responses occurred on a lever where responding initially had no scheduled consequence or, subsequently, when a 3-minute fixed-interval shock-presentation schedule was concurrently arranged for lever responses. Appropriate rates and patterns of lever responding developed and were later maintained under the fixed-interval 3-minute shock-presentation schedule alone when the chain and shock-postponement schedule were removed. When both the shock-postponement and shock-presentation schedules were again simultaneously in effect, steady rates of chain pulling were maintained by the shock-postponement schedule and positively accelerated rates and patterns were maintained on the lever by the shock-presentation schedule. Response rates under both schedules were directly related to shock intensity. A history of exposure to a shock-postponement schedule, even though with a topographically different response and manipulandum, was sufficient for the development and eventual maintenance of responding by the presentation of shock. Further, differential performances can be maintained simultaneously by the presentation and postponement of electric shock.     

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.     

Some effects of response-independent shocks after unsignaled avoidance conditioning in rats

After extended unsignaled avoidance training, the majority of rats continued frequent responding during long periods when only response-independent shocks were presented. Most responses were shock elicited, i.e., followed immediately after shock. Response rates were directly related to both the frequency and the intensity of response-independent shocks. Responding continued to approximately the same extent, whether shocks were presented at fixed or variable intervals. Few responses were directed toward a second lever in the test chamber, even when the lever previously associated with avoidance was removed. When avoidance was scheduled on the second lever, the rats learned to avoid by operating it. Meanwhile, responding on the first lever became infrequent. Subsequently, when only response-independent shocks were presented, almost all responses occurred upon the lever initially associated with avoidance. Responding that was elicited by response-independent shocks was suppressed by response-dependent (punishment)_shocks of the same intensity. When punishment was withdrawn, recovery of responding occurred. An explanation of the results based upon shock-elicited behaviors is preferred to one stressing unextinguished avoidance responding.     

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.     

Acquisition of cocaine self-administration with unsignaled delayed reinforcement in rhesus monkeys

Six experimentally naive rhesus monkeys produced 0.01 mg/kg/infusion cocaine by lever pressing under a tandem fixed-ratio 1 differential-reinforcement-of-other-behavior schedule. One lever press initiated an unsignaled 15- or 30-s delay culminating in cocaine delivery. Each press made during the delay reset the delay interval. With two exceptions, responding was acquired and maintained at higher rates than responding on a second (inoperative) lever. For the exceptions, a cancellation contingency was arranged in which each formerly inoperative-lever response reset the tandem schedule. This manipulation reduced presses on the inoperative lever. Subsequently, the consequences of responding on the two levers were reversed, and the monkeys again responded at higher rates on the operative lever. As a comparison, 3 additional experimentally naive monkeys received response-independent cocaine deliveries. Although lever pressing was observed, it extinguished and was subsequently reestablished under the tandem schedule. The results suggest that although response-reinforcer contiguity is not required for cocaine to acquire reinforcing functions, a response-reinforcer relation appears necessary.     

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.     

The relative aversiveness of signalled vs unsignalled avoidance

Subjects avoided shock by pressing on one lever under an unsignalled condition, but by pressing a separate lever they changed the condition to signalled avoidance for 1-min periods. Signalled avoidance periods were identified by a correlated stimulus. All eight subjects responded to change the unsignalled schedule to a signalled one. Once contact with signalled avoidance was made, subjects continued responding to remain in that condition. Other tests showed that changeover responding was greater when the correlated stimulus was presented without the signal than when the signal was presented without the correlated stimulus. An analysis based upon shock and shock-free periods is presented.     

Rate changes after unscheduled omission and presentation of reinforcement

Changes in response rate similar to frustration effects were studied in a two-lever situation. Responding on one lever on a fixed-interval schedule produced access to water for 5 sec and an exteroceptive stimulus. In the presence of this stimulus, responding on another lever on a fixed-interval schedule produced access to water for 5 sec and terminated the stimulus. Occasional omission of a previously scheduled reinforcer after responding on the first lever resulted consistently in increases in rate on the second lever during the immediately succeeding interval. In another procedure, occasional presentation of a previously unscheduled reinforcer after responding on the first lever resulted consistently in decreases in rate on the second lever during the immediately succeeding interval. Changes occurred after the first omissions or presentations and were about the same in magnitude as the procedure continued over several sessions. Typically, an increase or decrease in rate was maintained throughout an entire 100-sec interval. Changes in rate on the second lever of approximately the same magnitude also occurred when rate on the first lever was near-zero under a schedule that differentially reinforced behavior other than lever pressing.     

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.     

Predator videos and electric shock function as punishers for zebrafish (Danio rerio)

Zebrafish (Danio rerio) are a promising animal model for studying the effects of gene–environment interactions on behavior. Two experiments were conducted to assess punishment effects of presenting predator videos (Indian leaf fish; Nandus nandus) and electric shock on operant approach responses in zebrafish. In Experiment 1, the predator video and shock stimuli were presented upon a response maintained by a single variable‐interval schedule of food reinforcement in different groups of fish. In Experiment 2, the predator video and shock stimuli were presented upon one of two response alternatives maintain by concurrently available variable‐interval schedules of food reinforcement in different groups of fish. Responding decreased when the predator video and shock stimuli were presented relative to their absence in both experiments. Moreover, responding on an unpunished alternative did not reliably decrease in Experiment 2. These results indicate that the decrease in responding resulted from the punishment contingency rather than from elicited species‐specific defense responses or conditioned avoidance. Thus, the predator video and electric shock functioned as punishers of operant behavior for zebrafish. Identifying punishers for this species could lead to research on how gene–environment interactions influence individual differences in sensitivity to punishment.     

Effects of chlordiazepoxide and cocaine on concurrent food and avoidance-of-timeout schedules.

Five rats were trained on a concurrent schedule in which responses on one lever produced a food pellet on a random-interval 30-s schedule during 10 s of food availability associated with distinctive exteroceptive stimuli. Responses on another lever postponed for 20 s the presentation of a 50-s timeout, during which all stimuli were extinguished and the schedule contingencies on the food lever were suspended. The response rates maintained by the random-interval schedule exceeded those maintained by the avoidance contingency, but both provided a stable baseline to assess the behavioral effects of different drugs. Low doses of cocaine hydrochloride (1 and 3 mg/kg) did not affect food-reinforced responding or avoidance response rates. Intermediate doses (5.6, 10, and 13 mg/kg) produced a dose-dependent decrease in food-maintained and avoidance response rates, and both types of responding were virtually eliminated after administration of the highest doses (17 and 30 mg/kg) of cocaine. Low doses of chlordiazepoxide (1 and 3 mg/kg) increased food-maintained and avoidance response rates, and both rates decreased systematically after 10 and 30 mg/kg of this drug. The effects of cocaine and chlordiazepoxide on response rates maintained by avoidance of timeout from food presentation were unlike those reported when subjects responded to avoid shock presentation. The results of this experiment thus provide evidence to suggest that the effects of drug administration on avoidance behavior may be a function of the nature of the consequent event to be avoided.     

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.     

Behavior controlled by scheduled injections of cocaine in squirrel and rhesus monkeys.

Rates and patterns of key-press responding maintained under schedules in which responding resulted in intravenous injections of cocaine were studied in squirrel monkeys and rhesus monkeys. Each injection was followed by a 60- or 100-sec timeout period. Schedule-controlled behavior was obtained at appropriate cocaine doses in each species. Under FR 10 or FR 30 schedules, performance was characterized by high rates of responding (usually more than one response per second) in each ratio. Under FI 5-min schedules, performance was characterized by an initial pause, followed by acceleration of responding to a final rate that was maintained until the end of the interval. Under multiple fixed-ratio fixed-interval schedules, rates and patterns of responding appropriate to each schedule component were maintained. Responding seldom occurred during timeout periods under any schedule studied. At doses of cocaine above or below those that maintained characteristic schedule-controlled behavior, rates of responding were relatively low and patterns of responding were irregular. Characteristic fixed-interval responding was maintained over a wider range of cocaine doses than characteristic fixed-ratio responding. Complex patterns of responding controlled by discriminative stimuli under fixed-ratio or fixed-interval schedules can be maintained by cocaine injections in squirrel monkeys and rhesus monkeys.