A comparison of escape and avoidance conditioning in wild and domesticated rats

In the first of two experiments, three cotton rats (Sigmodon hispidus) and three albino rats were exposed to instrumental escape, unsignaled avoidance, and signaled avoidance, in that order. All subjects learned the escape procedure quickly, with the albino rats having generally shorter latencies, higher response rates, and requiring fewer sessions to reach the criterion. When the avoidance contingency was introduced, the cotton rats continued to respond almost entirely in the presence of the shock, whereas the albino rats responded in its absence, thus displaying effective avoidance behavior. Introduction of a pre-aversive stimulus did not improve the performance of the cotton rats. In the second experiment, five cotton rats and four albino rats were exposed to a free-operant (Sidman) avoidance procedure with a shock-shock interval of 3 sec and a response-shock interval of 20 sec. The cotton rats initiated responding at lower shock intensities than the albino rats, but their asymptotic avoidance responding was far less effective.     

Role of intertrial interval following an escape or avoidance response in bar-press avoidance

In three experiments, rats learned bar-press avoidance as a function of the intertrial interval following an escape or an avoidance. The general hypothesis is that the length of these intervals affects bar-press avoidance differentially, depending on whether an avoidance follows an escape (avoidance/escape) or an avoidance (avoidance/avoidance) on the previous trial. Specifically, it is proposed that short escape ITIs will facilitate avoidance/escape and avoidance/avoidance, while avoidance ITIs will have no effect on avoidance responding. The results tend to support this hypothesis. The shorter the intertrial interval following an escape, the higher the probability of both measures: avoidance/escape and avoidance/avoidance. No effect of avoidance interval was found on avoidance/avoidance. Unpredictedly, however, it was found that in comparison to a very short intertrial interval following an avoidance (0.5 sec), relatively long intervals (5 and 45 sec) facilitate avoidance/escape. These results were interpreted as mainly reflecting nonassociative factors such as shock-produced activity.     

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.     

Effects of reinforcement magnitude and ratio values on behaviour maintained by a cyclic ratio schedule of reinforcement

In Experiments 1 and 2, lever pressing by rats was reinforced on a cyclic ratio schedule of food reinforcement, comprising a repeated sequence of fixed-ratio component schedules. Reinforcement magnitude was varied, on occasional sessions in Experiment 1 and across blocks of sessions in Experiment 2, from one to two or three 45-mg food pellets. In the one-pellet condition, post-reinforcement pauses increased with component schedule value. At higher magnitudes, post-reinforcement pauses increased, and overall response rates declined. Response rate on component schedules was a decreasing linear function of the obtained rate of reinforcement in all conditions. Plotted against component schedule value, response rate increased exponentially to an asymptote that decreased when reinforcement magnitude increased. These findings are consistent with regulatory accounts of food reinforced behaviour. In Experiment 3, rats were trained under a cyclic ratio schedule comprising fixed-ratio components including higher values, and some inverted U-shaped response functions were obtained. Those rats that did not showthis relationship were trained on cyclic ratios with even higher values, and all showed inverted U-shaped response functions. This suggests that behaviour on cyclic ratio schedules can reflect activating of reinforcement as well as the satiating effects seen in Experiments 1 and 2.     

Stimulus control of responding during a fixed-interval reinforcement schedule

During training sessions, pecks by pigeons on a response key illuminated by a vertical line of white light resulted in reinforcement and an ensuing blackout according to a fixed-interval schedule. Training sessions were followed by dimensional stimulus control test sessions during which the orientation of the line present throughout the fixed interval was varied. Inverted U-shaped (excitatory) gradients of responding, with maximum responding occurring in the presence of the vertical line, were observed during the terminal part of the fixed interval. U-shaped (inhibitory) gradients of responding, with minimum responding occurring in the presence of the vertical line, were observed during the early part of the fixed interval when the preceding interval had terminated with reinforcement and blackout but not when the preceding interval had terminated with blackout only. These results suggest that the dimensional control by the stimulus present throughout the fixed interval is of a conditional variety. Whether the fixed-interval stimulus exerts inhibitory or excitatory dimensional control depends upon the presence and absence, respectively, of stimuli associated with reinforcement.     

Forgetting of an operant response: Physostigmine-produced increases in escape latency in rats as a function of time of injection

Latency of a fixed ratio (FR) 3 escape response in rats was found to be a U-shaped function of the interval between training and injection of the anticholinesterase drug physostigmine, for intervals from 30 min. to 5 days between training and injection. An increase in FR 3 escape latency was found at 28 days. FR 1 escape groups produced a latency curve of a shape similar to that of the FR 3 group. These data confirm the results of earlier experiments using a different training procedure, and a different response measure. These results are consistent with the theory that the physiological correlate of rat memory lies in synaptic change.     

Effects of parafascicular excitotoxic lesions on two-way active avoidance and odor-discrimination

To investigate whether the parafascicular (PF) nucleus of the thalamus is involved in different learning and memory tasks, two experiments were carried out in adult male Wistar rats that were submitted to pre-training bilateral N-methyl-d-aspartate PF infusions (0.15M, pH 7.4; 1.2 microl/side, 0.2 microl/min). In Experiment 1, we evaluated the effects of PF lesions in two identical 30-trial training sessions, separated by a 24-h interval, of a two-way active avoidance conditioning. PF-lesioned rats exhibited impaired performance in both sessions, measured by number of avoidance responses. In Experiment 2, the effects of PF lesions were assessed in a training session (5 trials) and a 24-h retention test (2 retention trials and 2 relearning trials) of an odor-discrimination task. PF lesions did not significantly disrupt the acquisition or the first retention trial, which was not rewarded. However, lesioned animals' performance was clearly affected in subsequent trials, following the introduction of the single non-rewarded trial. Current data are discussed considering evidence that lesions of the PF nucleus affect learning and memory functions mediated by anatomically related areas of the frontal cortex and striatum.     

Cardiovascular responses to avoidance conditioning in the dog: Effects of alpha adrenergic blockade

Laboratory dogs were trained to press a response panel to postpone shocks during daily one-hour avoidance conditioning periods. Each dog was also confined in the experimental environment for 5 hours prior to the avoidance periods. Blood pressure and heart rate were monitored continuously during these experiments from chronically indwelling arterial catheters. Extended training resulted in the emergence of a cardiovascular response pattern during the pre-avoidance interval characterized by gradual increases in blood pressure together with decreases in heart rate. Elevations in both blood pressure and heart rate were sustained during the avoidance periods. During sessions in which alpha adrenergic activity was suppressed by phenoxybenzamine, absolute levels of blond pressure were found to be lower than during control (non-drug) sessions, but a progressive rise in blood pressure continued to be observed during pre-avoidance. These results suggest that sustained cardiovascular responses during avoidance periods are associated with activation of the sympathetic nervous system, but that the gradual rise in blood pressure during pre-avoidance is due to other factors.     

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.     

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.     

The effects of aversive conditioned stimuli on the timing of unsignalled avoidance responding in rats

Rats received unsignalled avoidance training in a shuttlebox. After acquisition, classical conditioning sessions were administered. The Excitatory Group received signals paired with shock, the Inhibitory Group received signals explicitly unpaired with shock, while the Random Control Group received signals and shocks randomly in time. When subsequently superimposed on the avoidance baseline, the excitatory signal increased response rate, the inhibitory signal decreased rate and the random signal had no effect compared to signal-only control data. Unsignalled avoidance generates temporal gradients of responding; the major purpose of this study was to determine how these rate changes were reflected in the baseline temporal behavior. Very clear and simple summation rules emerged: In the presence of the excitatory signal, rats acted as if they were ahead of their actual location in the response-shock interval regardless of where the signal occurred; the inhibitory signal had the opposite effect while the random signal did not change the temporal behavior relative to signal-only control data. These results were interpreted within a motivational framework.     

The discriminative control of free-operant avoidance despite exposure to shock during the stimulus correlated with nonreinforcement

Four rats were trained in darkness on a free-operant avoidance procedure in which shocks occurred randomly, but lever presses could reduce their frequency. Discrimination training followed, during which responses in light continued to reduce shock frequency, but responses in darkness had no effect. During each cycle, the light period was 4 min, while darkness lasted only until a 20-sec interval had elapsed without a response. This no-response requirement was increased to 40 sec for three animals and eventually to 60 sec for two of them. Discriminative control developed, despite a greater shock density in the dark, with response rate and number of responses per shock maintained or increasing during light and decreasing to very low values in darkness. Two animals were later exposed to a procedure in which shock density was unaffected by responding either in light or darkness. A 60-sec no-response requirement was continued in the dark. Discriminative control persisted through 42 sessions for one animal and required 45 sessions to approach extinction for the other animal. The role of the light as a potential conditioned reinforcer of other behavior in the dark was implicated in the development and persistence of discriminative control. These data support shock-frequency reduction as reinforcement for avoidance behavior.     

Effects of food deprivation on the performance of a two-turn task in a temporal circular maze

Ten hooded rats were trained to perform a two-turn task in a temporal circular maze. The effects of levels of food deprivation (2, 26, 50, 74, 98, 122 hr.) on performance were evaluated. It was found that the correct responses were not affected, but both running time and competing behaviour decreased significantly with increase in hours of food deprivation. In a second experiment 8 rats were trained in a similar way and after attaining a stable performance, were run for a further 4 sessions at 24 hr. deprivation, and then at 72 hr. deprivation. The results precluded the possibility that the results obtained in Experiment I were due to practice effects, and support the findings on the effects of deprivation.     

Active role of the shock-avoidance contingency in shuttlebox-avoidance learning in Fischer344 rats

Different groups of Fischer344 rats acquired and maintained shuttlebox-avoidance responses under one, two, all three, or none of the shock-avoidance, warning-signal (WS) termination, and shock-escape contingencies. The intertrial interval was fixed at either 30 or 60 s. Removal of the shock-avoidance contingency led to a marked disruption of avoidance behavior at both intervals. This result was not produced by a selective punishment effect of the avoidance response by an inevitable shock. Thus, the shock-avoidance contigency provided particularly effective control over the avoidance learning. The WS termination contingency appeared to have little overall effect at the 60-s intertrial interval. It is suggested that response-contingent stimulus change served only as a discriminative cue for the absence of a shock.     

Punishment and the potential for negative reinforcement with histamine injection

The present study examined punishment of responding with histamine injection, and its potential to generate avoidance of punishment. Sprague–Dawley rats were trained under concurrent schedules in which responses on one lever (the punishment lever) produced food under a variable‐interval schedule, and under some conditions intermittent injections of histamine, which suppressed behavior. Responses on a second (avoidance) lever prevented histamine injections scheduled on the punishment lever. After stabilization of punished responding, a variable‐interval 15‐s schedule of cancellation of histamine (avoidance) was added for responding on the second/avoidance lever, without subsequent acquisition of responding on that lever. Progressive decreases in the length of the punishment variable‐interval schedule increased suppression on the punishment lever without increases in response rates on the avoidance lever. Exchanging contingencies on the levers ensured that response rates on the avoidance lever were sufficiently high to decrease the histamine injection frequency; nonetheless response rates on the avoidance lever decreased over subsequent sessions. Under no condition was responding maintained on the avoidance lever despite continued punishing effectiveness of histamine throughout. The present results suggest that avoidance conditioning is not a necessary condition for effective punishment, and confirm the importance of empirical rather than presumed categorization of behavioral effects of stimulus events.     

The role of the cs and avoidance period duration in discriminated avoidance conditioning

In an experiment investigating the effect of CS duration on discriminated bar-press conditioning, subjects were assigned to one warning period duration (1.5, 5.0 or 15.0 sec.) on the first day of avoidance training, and to one of the three durations on the second day of training. On each day avoidance behaviour was greatly influenced by the duration of the CS (warning) period, but the duration on day I had no effect on the second day's avoidance performance. Groups receiving no CS during training, although provided an avoidance contingency, showed little conditioning, but produced highly significant amounts of intertrial responding. In a second experiment, subjects receiving CS on day I were shifted to no CS on day II. Avoidance performance on day II was not significantly different from the day II performance of subjects in Experiment I having two training sessions with the CS present or absent on both days.     

Tolerance of pain as a measure of fear

Fear thresholds were measured in four experiments by exposing rats to electric shock in order to determine the maximal intensity rats would tolerate rather than enter a fear-arousing box and/or stop freezing. Increasing fear raised these thresholds. They were greater for rats having to escape shock to a fear-arousing box than for rats having to escape shock and fear to a neutral box. The forgetting functions for the latter two groups differed: the first group yielded a monotonic decay function, whereas the second group yielded an inverted U-shaped function. These thresholds decreased as a function of an avoidance learning procedure. Rats that had to escape shock to a fear arousing box did not do so immediately, although they had stopped freezing. An avoidance-avoidance conflict explanation for immobility was not found to be valid. A theoretical formulation based on the following two hypotheses was suggested to explain these results: the fear-aroused freezing (immobility) is an unlearned response; finding a way to escape the source of fear starts another unlearned response, withdrawal.     

Warmup in avoidance as a function of time since prior training

On avoidance procedures, rats and pigeons typically show warmup effects, characterized by improving performance within sessions and loss of the improvement (“warmup decrement”) between sessions. Between-session losses were examined by varying the time between periods of avoidance training. In one experiment, rats lived fulltime in conditioning chambers while intermission intervals were varied. In a second experiment, the animals lived in home cages between sessions; timeout intervals were introduced at midession, producing recurrence of warmup in the second half-session. In both experiments, the warmup decrements increased substantially as the timeout or intersession intervals were increased from zero to 30 minutes. With intervals of 60 or 120 minutes, the decrements approached or exceeded those obtained with intervals of a day or more. When avoidance was interposed between appetitive sessions, the appetitive responding was disrupted, but this seemed unrelated to the warmup or to the proficiency of avoidance. The warmup in avoidance shares characteristics with transient punishment effects, with the Kamin effect, and with habituation phenomena, but it is premature to assume that they reflect common processes.     

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.