Persistent shock-elicited responding engendered by a negative-reinforcement procedure

A procedure in which responses reduced intermittently presented electric shocks to one quarter of their originally scheduled intensity, effectively engendered and maintained lever pressing in hooded rats. This contingency also markedly increased the response rates of rats initially trained under an unsignaled avoidance procedure. The responding of all animals extinguished rapidly when shock was withdrawn. Subsequently, it was discovered that high response rates could be maintained solely through presentation of shocks that were not affected by responses. Variations in the interval between shocks and changes in shock intensity over a wide range did not attenuate responding. Terminal performance was characterized by a consistent pattern of shock-elicited responses. Responses were also elicited by a tone following repeated tone-shock pairings. Finally, responding that was maintained by response-independent shocks was quickly suppressed by response-contingent shocks of the same intensity.     

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.     

Acceleration and suppression of rats' responding to avoid foot shock and tail shock

Signalled response-independent shocks were superimposed on rats' wheel-turn responding to avoid shock administered to their feet through a grid floor or to their tails through fixed electrodes. In Experiment I, a tone paired with response-independent foot shock increased responding in three of four rats; a tone paired with tail shock increased responding in only one of four rats and suppressed responding in two rats. In Experiment II, a tone presented randomly with respect to response-independent shock had no reliable effect on responding to avoid foot shock or tail shock. In Experiment III, tail shock and foot shock were compared in a within-subject design while the temporal pattern of responding during conditioned stimuli was recorded. Responding during the conditioned stimulus preceding foot shock was characterized by initial suppression of responding at tone onset, followed by increased responding just before response-independent shock. Responding was suppressed throughout the conditioned stimulus preceding tail shock. Foot shock elicited bursts of responding, but tail shock did not.     

Responding in the cat maintained under response-independent electric shock and response-produced electric shock

Key-pressing responses in the cat were maintained under conditions in which brief electric shock was first postponed by responses (avoidance), then periodically presented independently of responses, and finally produced by responses on a fixed-interval schedule of 15 min (FI 15-min). A steady rate of responding occurred under shock avoidance and under response-independent shock; positively accelerated responding was engendered by the FI 15-min schedule. A second experiment studied responding under second-order schedules composed of three FI 5-min components. Responding was suppressed when a stimulus was presented briefly at completion of each FI 5-min component and a shock followed the brief stimulus at completion of the third component. Responding was maintained when each of the first two components was completed either with or without presentation of a brief stimulus and a shock alone was presented at completion of the third FI 5-min component.     

The effect of rate of delivery of response-independent shocks upon avoidance responding

Two hooded rats were trained to bar-press to avoid electric shock on a continuous avoidance schedule with response-shock and shock-shock intervals equal. The rate of delivery of response-independent shocks superimposed on this schedule was varied. The response-independent shocks led to generally higher response rates but, with responses during shock omitted, the rates decreased as the response-independent shock rate was increased. The actual shock rate received by the subjects was linearly related to the maximum potential shock rate. There was an increasing, negatively accelerated function between percentage avoidance and response rate, but there was no consistent relation between the number of shocks avoided and response rate. Response rate decreased as the potential shock rate increased, but responding was maintained even when as few as 15% of the shocks could be avoided.     

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.     

Reduction of intraspecies aggression in rats by positive reinforcement of incompatible behaviors

Fighting responses were elicited by response-independent shocks delivered to pairs of rats. Food pellets were presented following different non-fighting responses to shock: some pairs of rats received pellets dependent upon a specific non-fighting response to shock; others received pellets dependent upon any non-fighting response to shock; and control pairs never received pellets. The mean probability of an elicited aggressive response to shock was reduced to 0.2 by food reinforcement for a specific non-fighting response, and to 0.5 by food reinforcement for any kind of non-fighting response. These values contrasted with the 0.8 probability of elicited aggression when pairs of rats received no food reinforcement. Consistent findings were obtained when treatment conditions were changed for individual pairs of rats.     

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”.     

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.     

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.     

Discrimination of response-contingent and response-independent shock by rats: effects of chlordiazepoxide HCL and sodium amylobarbitone

Rats, trained to press a lever for sucrose reward on a random interval (RI) schedule, were presented while lever-pressing with two stimuli, each associated with a different schedule of shock delivery: in the presence of one stimulus (S_e), shock occurred on an RI schedule irrespective of the rat's behaviour; in the presence of the other (S_p) shocks were programmed by the same schedule but delivered only when the rat pressed the lever. Both stimuli suppressed lever-pressing. In addition, the rats developed significantly different response rates in the two stimuli, thus demonstrating a discrimination between response-contingent and response-independent shock. Group data showed faster responding in S_e than in S_p, supporting the view that response-contingent shock produces greater suppression than response-independent shock. Individual animal analyses, however, demonstrated that this was the case in the majority of animals, but not in all. Response suppression was alleviated by amylobarbitone sodium (15 mg/kg) or chlordiazepoxide HCI (5 mg/kg); the latter drug alleviated suppression significantly more in S_p than S_e and eliminated the difference between the response rates controlled by the two stimuli.     

The effects of stimuli followed by response-independent shock on shock-avoidance behavior

Four rhesus monkeys were trained on a non-discriminated shock-avoidance schedule (baseline). Stimuli followed by response-independent shock were then presented with the avoidance baseline no longer in effect. The main portion of the experiment consisted of superimposing (independently of responding) the stimuli followed by response-independent shock on the avoidance baseline. Different temporal values of stimulus duration and delay of shock (produced by an avoidance response) were presented successively, using each subject as his own control. When the stimulus duration was short or the delay of shock was long, so that avoidance rate during the stimulus could assume any value without resulting in baseline (avoidable) shocks during the stimulus, a lowered or "suppressed" rate of responding developed during the stimulus. When the stimulus duration was long or the delay of shock was brief, so that avoidable shocks resulted from a response decrement during the stimulus, high or "facilitated" rates of responding developed for a large proportion of the time that the stimulus was present.     

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.     

Lever attacking and pressing as a function of conditioning and extinguishing a lever-press avoidance response in rats

Six experimental rats were conditioned to press one of two available levers to avoid shock. The levers registered bites as well as presses. For four of these rats, shock was contingent on lever bites when a specified time period had elapsed after the previous shock. An extinction period, in which only periodic noncontingent shocks were presented, followed avoidance training. Six yoked-control rats received the same sequence of shocks as did the corresponding experimental rats in both the conditioning and extinction phases. All six experimental rats repeatedly bit the avoidance lever. Four bit it more than the nonavoidance lever during conditioning, and five bit it more during extinction. Five of the six experimental rats consistently bit the levers many more times during each session than did their respective control rats, suggesting that avoidance conditioning facilitated lever biting. Rates of lever biting and pressing by all of the experimental rats and by some of the control rats were highest immediately following shock throughout both phases. During later portions of the intervals following shock, characteristic effects of conditioning and extinction were observed. This finding suggests that extinction of avoidance behavior by unavoidable shock presentations can be demonstrated more readily when shock-elicited responding is extricated from the data.     

Ratio responding as a function of concurrent avoidance schedules, yoked shocks, and ratio value

Fixed-ratio food-reinforced responding in rats was studied alone and with concurrent shock avoidance or with concurrent response-independent shocks matched to those that occurred in the avoidance condition. Under each condition, fixed-ratio size was increased over successive daily sessions. Fixed-ratio response rate generally passed through a maximum as a function of fixed-ratio size. Decreased fixed-ratio responding at values beyond the maximum occurred when (1) the time to complete a fixed ratio approximated the response-shock interval of the avoidance schedule, (2) the shock rate increased, and/or (3) the ratio requirements were so high that ratio strain occurred. Avoidance rates decreased slightly as fixed-ratio size increased.     

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.     

What is both necessary and sufficient to maintain avoidance responding in the shuttle box?

To learn what maintains the frequency of shuttle box avoidance responses, male rats from the Berkeley S₁ strain, after 200 trials of standard discriminative avoidance training, were given 100 additional trials under one of four different conditions. Responding at the maximum rate was maintained when animals performed under the training conditions or when responses continued to terminate the warning signal immediately, even though shock was never given for failing to respond. In contrast, avoidance responding was reduced markedly if, and only if, trials were given in which the signal ceased to terminate immediately (i.e. it shut off either well before or well after a response). This decrement occurred even though avoidance responses continued to avert shock. Thus, under the conditions of this experiment prompt signal offset was both necessary and sufficient to maintain the occurrence of well-established shuttle box avoidance responses.     

The effects of escape conditioning and shock intensity on responding during inescapable shock

Eight albino rats, conditioned to press a lever to escape shock, continued to lever press during short inescapable shocks presented subsequently. The rate of this behavior was found to be higher for higher shock intensities regardless of the order in which shock values were presented. Relative to the immediately preceding escape rate, responding during inescapable shock was higher following conditioning at higher fixed-ratio escape requirements. Four subjects not conditioned to escape shock pressed the lever very infrequently during inescapable shock and showed little change with changes in shock intensity. The escape conditioning effects suggest that responding during inescapable shock is superstitious escape behavior. The effects of shock intensity on this behavior appear to be similar to reported effects of shock intensity on escape behavior.     

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.     

Lever attacking by rats during free-operant avoidance

Rats pressed a lever to avoid shock on a free-operant avoidance schedule. Some subjects were also exposed to extinction in which the response-shock contingency was eliminated while the shock-shock contingency remained in effect. A specially constructed lever was used that registered not only presses, but also biting attacks on the lever. Throughout various phases of the study, shocks often elicited lever biting as well as post-shock responding. The results suggested that shock-elicited attacks that are forceful enough to activate the operandum might account for some of the responding that occurs in experiments on free-operant avoidance behavior. In particular, shock-elicited operandum attacking might account for post-shock response bursting during free-operant avoidance and the extreme persistence of responding sometimes noted when shocks are delivered during the extinction of avoidance behavior. To the extent that this is true, these phenomena should not be characterized as operant behavior in interpreting the results of experiments on free operant avoidance.