Vicarious conditioned acceleration: successful observational learning of an aversive Pavlovian stimulus contingency

Five rats (observers) were trained to avoid unsignalled shocks in a shuttlebox and then habituated to brief light presentations. They were next confined on an observation platform while another rat (model) received light-shock pairings in the opposite compartment. The observers were exposed only to the sight and sound of the model during classical conditioning and were not shocked themselves. Test presentations of the light during subsequent avoidance sessions produced response rate increases (vicarious conditioned acceleration) comparable to those obtained in other studies where the avoidance animals were used in classical conditioning. Following sessions in which the model was not shocked after the light, the light presentations during avoidance eventually failed to elicit any response increases in the observers. When the model was again shocked, immediate recovery of avoidance acceleration occurred in the observers during the light.     

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 restoration of extinguished avoidance responses

Following extensive signaled shuttlebox avoidance training, Fischer 344 rats were given extinction trials until they reliably stopped responding and, thereafter, they received occasional classical CS-UCS pairings during the course of restoration test trials in which shock was absent. Experiment I demonstrated that the classical contingency was both necessary and sufficient for the complete recovery of such behavior. Experiment II revealed that the degree of recovery was not differentially affected by the frequency of such CS-UCS pairings. However, in Experiment III recovery in the presence of this contingency was shown to depend on the number of previous extinction trials, although even after a large number of such trials, a considerable degree of recovery occurred. These findings demonstrated the central role of the classical CS-UCS contingency in restoring a previously extinguished avoidance response.     

On the independence of stimulus evocation of fear and fear evocation of responses

Dogs in an experimental group were first trained in a shuttlebox on a free-operant avoidance response; then a second, discriminative avoidance response was trained in a second apparatus to a tone. Following criterion, the second avoidance response was “extinguished” through unsignaled. response-independent US presentations. Finally, the tone was presented in the shuttlebox as a test for residual fear. The tone produced as much facilitation of the free-operant avoidance responding as it did in two control groups; one control had not had the second avoidance response extinguished, while the other had received only Pavlovian conditioning in the second apparatus. Thus, the elimination of a discriminative avoidance response to a CS does not impair the CS's ability to control a second independent avoidance response. This result reflects and confirms the separability and independence of the CS-fear and fearavoidance response links hypothesized by two-process theories.     

A semiautomatic method for adding behavioral data to videotape records in combination with the Noldus Observer system

The Observer 3.0 (Noldus, 1991) was used as the base for a time-saving fine analysis of animal behavior. A PC controlled a shuttlebox and a shuttlebox control unit. Learning behavior was recorded on videotapes by an S-VHS camcorder with vertical interval time code (VITC), which was used to generate an observational data file (ODF). The observational system was enhanced by automatically recorded status signals from the PC as event markers on the soundtrack of the videotape. These signals were used to subdivide a training session into numbered trials during generation of the ODF. They were later used for a PC-aided fine analysis of behavioral events, such as attention responses (AR) and orienting responses (OR) during shuttlebox learning. In this way, a large number of both automatically inserted PC event markers and manually inserted, key-defined behavioral events were registered and analyzed effectively in a semiautomatic fashion. The system is illustrated with an analysis of shuttlebox avoidance data from gerbils (Meriones unguiculatus).     

Differences in learning between hyperprolinemic mice and their congenic controls

These experiments expanded earlier work on hyperprolinemic mice which showed learning deficits. The following behavioral tasks were used: step-through, passive avoidance; T-maze acquisition; shuttlebox acquisition, and radial-arm maze. Mouse species included PRO/Re-bb (genetically hyperprolinemic mice) and PRO/Re-aa (congenic nonhyperprolinemic controls) obtained from the Jackson Breeding Laboratories. Hyperprolinemic mice were impaired in acquiring T-maze and shuttlebox footshock avoidance behavior. One-trial passive avoidance behavior did not clearly differentiate between the groups. Radial maze performance was poor in both groups due possibly to observed acrophobia and lack of exploratory behavior. The results of this study combined with previously published work suggest that high-brain proline in conjunction with other amino acid changes account for the learning deficits.     

Prior exposure to inescapable electric shock in rats affects extinction behavior after the successful acquisition of an escape response

In Expt 1, rats exposed to 64 inescapable electric shocks in a restrainer or merely restrained were later given either 0, 5, 15 or 30 escape/avoidance training trials with a two-way shuttlebox procedure that does not lead to interference with escape acquisition due to prior exposure to inescapable shock. After escape training all rats were given an escape/avoidance extinction procedure in which shock was inescapable. The rats which had received prior exposure to inescapable shock responded less often and with longer latencies in extinction than did the restrained rats. Experiment 2 demonstrated that this effect is caused by the inescapability of the initial shock treatment. These results were explained in terms of (a) associative interference which minimized the effect of shuttlebox escape training for the preshocked subjects, and (b) a stronger tendency to recognize the presence of an inescapable shock situation during extinction for the preshocked subjects. The relationship between these results and previous work demonstrating that exposure to the escape contingency mitigates the effects of inescapable shock exposure was also discussed.     

Signaled avoidance in the eye withdrawal reflex of the green crab

Learning in a signaled avoidance procedure was studied in the eye withdrawal reflex of the green crab, Carcinus maenas. A puff of air to the eye, which causes eye retraction, was used as the unconditioned stimulus (US). A mild vibration on the carapace, which has no effect on untrained animals, was used as a warning (conditioned) stimulus (CS). Eye withdrawal during the CS led to the omission of the otherwise scheduled US. Acquisition was rapid, reaching about 75% avoidance after 30 trials. Extinction occurred slowly over the course of 40 CS-only trials. Yoked controls did not perform as well. The behavior of experimental animals in the avoidance procedure was found to be essentially identical to the performance of animals subjected to a classical conditioning paradigm in which CS responses had no effect on US presentation. Additional groups of animals were subjected to experiments in which (a) avoidance conditioning (60 trials) was followed by classical conditioning (40 trials) or (b) classical conditioning was followed by avoidance. The behavior of these groups was, again, essentially identical. The results suggest that there may be an underlying Pavlovian mechanism for the learned response, although the contribution of an operant process is not excluded. The results expand the range of invertebrate animals in which fundamental conditioning phenomena can be demonstrated, and may provide a neuronal model for learning in a signaled avoidance procedure.     

Therapy for helpless monkeys

During the course of pilot studies and two formal experiments examining the learned-helplessness phenomenon in rhesus monkeys, 5 subjects failed to escape in a shuttlebox following earlier experience with aversive stimulation in primate-restraining chairs. The present report details a therapeutic manipulation designed to reverse these subjects' maladaptive behavior in the shuttlebox. Introduction of a different fear stimulus (a net previously used to restrain the animals) was found to be effective in inducing shuttlebox escape and avoidance learning. The implications of the present findings for an understanding of the learned-helplessness phenomenon and their relevance to therapy for human depression are discussed.     

Note on the invariance of response latency in shuttlebox avoidance learning

Rats trained to avoid shock by running in a shuttlebox showed an orderly increase over trials in the probability of avoidance but no corresponding change over trials in the latency of the avoidance response. Both responding in the presence of shock (escapes) and responding prior to shock onset (avoidances) appeared to be stationary in time. Latencies were found to vary with different experimental conditions but in every case to be invariant over trials.     

Role of fear in mediating shuttle escape learning deficit produced by inescapable shock

The relation between the shuttlebox escape deficit produced by prior inescapable shock (IS) and fear during shuttlebox testing as assessed by freezing was investigated in rats. IS rats learned to escape poorly and were more fearful than either escapably shocked subjects or controls, both before and after receiving shock in the shuttlebox. However, fear and poor escape performance did not covary with the manipulation of variables designed to modulate the amount of fear and the occurrence of the escape deficit. A 72-hr interval between IS and testing eliminated the escape deficit but did not reduce preshock freezing. Diazepam before testing reduced both preshock and postshock fear in the shuttlebox but had no effect on the escape deficit. Naltrexone had no effect on fear but eliminated the escape deficit. This independence of outcomes suggests that the shuttlebox escape deficit is not caused by high levels of fear in IS subjects.     

Pavlovian conditioned inhibition of fear during shuttlebox avoidance behavior

Three experiments explored the development and function of conditioned inhibition of fear during the acquisition and maintenance of shuttlebox avoidance behavior. The development of inhibition to an exteroceptive feedback stimulus was found to be a function of the number of successive avoidance responses to which the animal had been trained and of the duration of the intertrial interval, a parameter shown also to affect the rate of acquisition of avoidance learning. Master animals who learned the instrumental avoidance response, and yoked animals who did not, showed equivalent inhibitory fear conditioning in each experiment. The results of one experiment suggest that conditioned inhibition plays no important role in “protecting” fear conditioned to the discrete warning signal during avoidance maintenance. These data indicate that feedback stimuli develop their inhibitory properties by a Pavlovian process and that certain aspects of their function may, therefore, be readily understood within the framework of mediational two-process learning theory.     

Framing of Decisions: Effect on Active and Passive Risk Avoidance

Decision makers intending to avoid risk in a decision situation can choose a less risky alternative (passive risk avoidance) or intervene actively in an alternative applying a risk‐defusing action (active risk avoidance). In Experiment 1 (64 participants), we compared active and passive risk defusing in two framing conditions. In the negative frame, in the uncertain alternative, a change to the worse was possible; in the positive frame, a change to an improvement was possible. Each participant decided in both framing conditions. As expected, active risk avoidance behavior for preventing a negative outcome (i.e., in the negative frame) was more likely than for promoting a positive one (i.e., in the positive frame). If decision makers did not or could not actively defuse the risk, they chose in correspondence to the classical pattern: risk avoidance in the positive frame and risk seeking in the negative one. We replicated the latter result in a second experiment (32 participants). The classical framing pattern in passive risk avoidance in both experiments is remarkable, because participants were not presented or did not search for exact probabilities. Copyright © 2014 John Wiley & Sons, Ltd.     

Freezing as an avoidance response: Another look at the operant-respondent distinction

Groups of rats were trained with shock either contingent on freezing (punishment procedure) or contingent on not freezing (avoidance procedure). Although the different contingencies produced different levels of freezing behavior, these levels were attained immediately rather than over a number of trials. This result, together with the results of control rats, suggest that while freezing can be controlled by both punishment and avoidance procedures, in both cases the effects on freezing are due to elicitation rather than learning.     

Can a shuttlebox computer analog be used to condition human avoidance behavior?

Freedman (1990) introduced a computer analog of the shuttlebox paradigm and presented results to show that this analog could be used for studying human avoidance conditioning. In the present study, a first experimental phase was conducted to test how the fact that the tone was considered aversive in the instructions and the intensity of this tone affect avoidance behavior. In a transfer-of-control test phase, it was tested as to whether the warning stimulus presented as a cue to the aversive stimulus had acquired either aversive or informative quality. An effect of instructions was observed for both levels of the auditive stimulus, and a stimulus effect was found for those groups that were given the instructions that described the tone as aversive. In the case of subjects who in the first phase achieved a certain learning criterion, it was recorded how often they in the second phase selected a condition in which the warning signal of the first phase was not presented. No transfer of control was observed. Thus, no positive evidence was found indicating that Freedman’s computer analog could be used for studying human avoidance conditioning.     

Dissociative effects of different types and amounts of nonreinforced CS exposure on avoidance extinction and the CER

Two experiments examined the effectiveness of two amounts of flooding or response-prevention on hastening avoidance response extinction and on reducing CS-produced suppression of bar-pressing for food. In Experiment 1, 20 and 30 flooding trials were both shown to be effective in hastening the extinction of a well-learned shuttlebox avoidance response. In Experiment 2, rats trained under comparable conditions to those in Experiment 1 were tested following flooding for the CER in a different apparatus. The results indicated that 30, but not 20, flooding trials were sufficient to reduce the CER. In each experiment the results of additional control groups equated with the flooded groups for nonreinforced CS exposure also revealed a dissociation between the effectiveness of this CS time control procedure in hastening avoidance response extinction and in reducing the CER. Further comparisons showed that although 30 flooding trials did reduce the CER, the same total duration of nonreinforced CS Exposure in the form of avoidance extinction trials did not. Thus the context in which CS exposure occurs may affect the dynamics of extinction of the CER. The experiments are discussed in the broader context of dissociation of various indices of fear in humans.     

Avoidance extinction in goldfish

Independent groups of goldfish trained to avoid shock in a shuttlebox situation were presented with several extinction procedures in which the relationships between the conditioned stimulus and shock were altered and/or response contingencies removed. Random shock presentations, equivalent to the number of shocks received during avoidance acquisition, resulted in response decrements similar to those obtained when the conditioned stimulus was presented alone. Pairing the conditioned stimulus with shock on every trial, however, served to maintain response levels. When response-contingent punishment was superimposed upon these Pavlovian pairings, performance was facilitated slightly although punishment alone resulted in somewhat faster response reduction than that produced by exposure only to the conditioned stimulus. Extinction of avoidance responding produced by exposure to the conditioned stimulus alone was dependent on the total duration of exposure and independent of both number of stimulus onsets and response prevention. These experiments demonstrated that, in general, the procedures used to reduce avoidance responding in rats were equally effective for goldfish, with one exception: the introduction of a Pavlovian contingency following avoidance acquisition, making the previously avoidable shock unavoidable, maintained response probabilities near previously established levels.     

Dissociation between conditioned emotional response and extended avoidance performance

The present experiments examined the dissociation between the strength of a shuttlebox avoidance response (AR) and one index of fear of the avoidance CS. Avoidance response strength was indexed by resistance to extinction of the AR and by changes in response latency, and fear of the CS was indexed by the conditioned emotional response (CER) technique. Experiments 1, 2A, and 3A all found that rats trained to a criterion of 27 consecutive avoidance responses (CARs) showed response strength comparable or superior to rats trained to a criterion of 9 CARs. Experiments 2B and 3A demonstrated that rats trained to 27 CARs showed less suppression of bar pressing during the avoidance CS (less CER) than rats trained to 9 CARs. Experiment 3A also found that, when extinguished in the shuttlebox to a moderate criterion (5 consecutive trials without a response) before CER testing, rats trained to 9 CARs showed some, although not complete, loss of CER, whereas rats trained to 27 CARs showed no loss of CER. In Experiment 3B rats that took 1 vs 2 days to reach a criterion of 27 CARs were compared for their AR strength and for their CER. Although rats taking 2 days to reach criterion showed somewhat greater resistance to extinction of the AR than rats reaching criterion in 1 day, this variable had no apparent effect on the CER. Implications of the present results for current theories of avoidance learning are discussed.     

Analgesic and opioid involvement in the shock-elicited activity and escape deficits produced by inescapable shock

Three experiments examined the involvement of analgesic processes and endogenous opioids in the production of the shuttlebox escape acquisition and unconditioned activity deficits which follow exposure to inescapable shock. Experiment 1 found that the opiate antagonist naltrexone administered before the inescapable shock session interfered with the shuttlebox escape acquisition deficit which would normally follow. Experiment 2 found naltrexone to completely prevent the unconditioned activity deficit. The final experiment revealed that dexamethasone, a synthetic glucocorticoid which abolishes the analgesia produced by inescapable shock, reversed the activity deficit. These results indicate that endogenous opioids may be involved in the production of both the escape acquisition and activity deficits. They also suggest that the analgesia produced by these opioids may participate in the mediation of the activity deficit, even though analgesia is not involved in producing the shuttlebox acquisition deficit.     

The effects of overtraining on flooding of jump-up and shuttlebox avoidance responses

Two experiments are reported which test the effects of overtraining (2, 4 or 6 days) on the efficacy of flooding (response-prevention) in hastening the extinction of jump-up and two-way shuttlebox avoidance responses. In the jump-up box, overtraining reduces the effectiveness of flooding in two ways: Rats trained for 6 days and given a flooding treatment are more resistant to extinction than rats trained for only 2 or 4 days and given a flooding treatment. By contrast, rats trained for 6 days and given a control treatment are less resistant to extinction than control rats trained for 2 or 4 days. In the shuttlebox, overtraining neither reduces the effectiveness of flooding in hastening avoidance response extinction, nor reduces the resistance to extinction of control animals. Implications of these different effects of overtraining on jump-up and two-way shuttlebox responses are discussed.