Inescapable shock and food-competition dominance in rats

Inescapable electric shock disrupts escape-avoidance learning in another apparatus. This study demonstrates a deficit in a nonlearning task in which no aversive stimulus occurs. In Experiment 1, inescapable shock lowered rats’ dominance in a food-competition situation relative to restrained controls. In Experiment 2, inescapable shock lowered rats dominance in the same food-competition situation relative to a group that received the equivalent amount of escapable shock, demonstrating that the inescapability of the shock caused at least part of the decrement observed in Experiment 1. Experiment 3 does not find that inescapable shock caused a significant difference in food consumed or running time when the rats were tested alone, showing it unlikely that the dominance effects were caused by decreased hunger or reduced running following inescapable shock.     

Stimulus control of immunization against chronic learned helplessness

Two experiments investigated the effectiveness of multiple (five) sessions of signaled eseapable-shock pretraining in preventing (immunizing against) the shack-escape impairment produced by an equal number of sessions of signaled inescapable shock. In Experiment 1, rats were exposed to 50 pairings per session of a white-noise stimulus with escapable shock during the immunization phase. Subsequently, they were exposed to 50 pairings per session of a different (houselight) stimulus with inescapable shock. Shock-escape performance in a shuttlebox test with constant illumination revealed no evidence of immunization relative to the performance of rats given five prior sessions of light-signaled inescapable shock only. Experiment 2 was identical in all respects to Experiment 1, except that both the escapable- and the inescapable-shock phases for animals in the immunization treatment group involved the same stimulus (houseüght) as a shock signal. Under these circumstances, the prior escapable-shock training significantly reduced the shuttle-box escape deficit engendered by chronic exposure to signaled inescapable shock; performance in the shuttle-box was not reliably different from that of rats exposed to signaled escapable shock alone. These findings suggest that, under chronic conditions, the development of stimulus control using Pavlovian conditioning procedures may serve to modulate the normally prophylactic influence on later shock-escape acquisition of serial exposure to escapable and inescapable shocks.     

Avoidance,classical, and pseudoconditioning as a function of species-specific defense reactions in high- and low-avoider rat strains

Rats of the Australian High Avoider (AHA) and Australian Low Avoider (ALA) strains and their reciprocal crosses were exposed to 50 trials of one of three shuttlebox procedures. The avoidance group received pairings of a tone and shock. If the animals shuttled during the tone, they avoided the shock. If they waited until the shock came on, they could then escape it. The classical group received pairings of the tone and a brief inescapable shock. If they shuttled during the tone, the tone ceased and they immediately received the shock. If they did not shuttle, they received the brief shock at the termination of the tone. The pseudoconditioning group received the tone and the shock explicitly unpaired. If they shuttled during either the tone or the shock, the stimulus was terminated. There was no acquisition of anticipatory responding under the pseudoconditioning procedure. All groups evidenced an increase in anticipatory responding over trials under the classical procedure. The AHAs acquired the response faster and reached a higher asymptote than did the ALAs. Performance of the two reciprocal crosses fell in between. A similar pattern was observed under the avoidance procedure, albeit at slightly higher response levels. Subsequent studies established that the AHAs acquired a one-way avoidance response quickly, but were impaired on a passive avoidance task, whereas the reverse was the case for the ALAs. The reciprocal crosses were proficient at both tasks. These results suggest that shuttlebox avoidance is largely accounted for by classical conditioning of the predominant defensive response. When that response is compatible with performance on the task, acquisition is rapid (AHAs), and when it is not, acquisition is slow (ALAs).     

Inescapable shock interferes with the acquisition of an appetitive operant

This article reports the reinforcer generality of the interference effect resulting from exposure to inescapable shock. In Experiment 1, rats that received inescapable shock showed weak interference with the acquisition of an appetitive operant compared to animals exposed either to escapable or no shock. In Experiment 2, the response-reinforcer contingency was degraded by introducing a 1-sec delay of reinforcement on the appetitive task. Inescapable shock produced much stronger interference with the acquisition of the operant response than in Experiment 1. The results demonstrate reinforcer generality of the debilitating effects produced by inescapable shock.     

Reward magnitude and shock variables (continuity and intensity) in shuttlebox-avoidance learning

In Experiment I, eight groups of rats (n = 20) were given shuttlebox-avoidance training. Two levels of shock (.3 and 1.6 mA) were combined factorially with two levels of reward (large and small) under both continuous and discontinuous (.75 sec on and 2.00 sec off) shock. Visual situational cues were absent after a shuttle response for the large-reward condition and present for the small-reward condition. Superior performance was obtained with weak rather than strong shock under both reward conditions and with large rather than small reward only under the weak-shock condition. Continuity of shock had no differential effect on performance. Experiment II allowed the conclusion that the reward effect was attributable to a reinforcement mechanism. The data were taken as support for the effective reinforcement theory, which emphasizes the importance in avoidance learning of fear conditioned to situational cues.     

Escape acquisition following inescapable shock in the rat

Male rats which had received approximately 21 min of pulsed, inescapable tail shock during a 6-h session in a wheel-turn chamber were markedly deficient in acquisition of an FR 2 crossing escape response in a shuttlebox when first tested 22 or 70 h later (Experiments 1 and 2). Rats which had received identical amounts and patterns of escapable/avoidable shock, however, were not deficient (Experiment 1). Preventing wheel-turn responses during the inescapable shocks prevented the occurrence of the subsequent escape deficit, whereas reducing the feedback provided for the first crossing response of the FR 2 requirement enhanced the deficit (Experiment 3). These data can be best explained by the learned helplessness hypothesis and indicate that the types of responses available and made during the inescapable shocks are more important than previously indicated.     

The nature of the initial coping response and the learned helplessness effect

The availability of an effective coping response has been shown to attenuate the deleterious behavioral and physiological consequences of inescapable electric shock. In the current study, two groups of rats could escape tailshock by turning a wheel. When short-latency responses that appeared to be elicited by shock onset were permitted to terminate shock, rats subsequently failed to learn to escape in a shuttlebox and did not differ from rats which received an equivalent amount of inescapable shock. However, when a relatively long-latency response was required and short-latency responses were not allowed to affect shock, rats subsequently readily learned to escape in the shuttlebox. The implications of these results for explanations of the manner in which prior exposure to shock influences subsequent escape learning were discussed.     

Differential response gradients to a Pavlovian safety signal following active avoidance training in young and adult rats

Following 300 training trials in two-way shuttle avoidance signaled by a tone (CS+), two groups each of weanling and adult rats were given Pavlovian discrimination training in which the CS+ was followed by inescapable shock, and a more intense tone (CS—) signaled no shock. An additional group at each age level received both tones paired randomly with shock or no shock. Subsequent generalization tests along the frequency dimension indicated that both pups and adults tested at the CS+ intensity showed similar gradients of frequency control. Gradients for the adults tested at the CS — intensity tended to be inverted, with least responding at CS—, a result not found in the young subjects. The results were considered in light of Pavlovian extradimensional influences on the control of avoidance behavior.     

A comparison of signaled vs. unsignaled free-operant avoidance in wild and domesticated rats

Three experiments compared the performance of domesticated hooded rats and nondomesticated black rats under signaled and unsignaled free-operant leverpress avoidance. There was no difference between groups in asymptotic shock rates under unsignaled avoidance; however, the black rats avoided much more successfully when the signal was present, while the hooded rats showed little or no improvement. When a longer signal was employed (10 vs. 5 sec), the effects were essentially the same. The black rats generally had higher response rates, and this difference was most pronounced in extinction, where the hooded rats made very few responses. The domesticated rats received a disproportionately large number of shocks early in the session (warm-up) under unsignaled avoidance, but this tendency was much less pronounced for the black rats. However, both groups showed appreciable amounts of warm-up during signalled avoidance. The findings are discussed in terms of differences in levels of activation/arousal between domesticated and nondomesticated animals.     

The effects of order of shock durations on helplessness in rats

Whereas rats exposed to a series of progressively decreasing shock durations show deficits in shuttle-escape performance 24 h later, the same number and intensity of shocks in the reverse (increasing) order of durations does not produce the “learned helplessness” effect (Balleine & Job, 1991). We conducted two experiments to establish the generality of this shock-duration order effect on other measures of distress and helplessness in rats. In Experiment 1, rats exposed to decreasing durations of inescapable shock showed reduced consumption of quinine-adulterated water (finickiness), whereas increasing durations produced no finickiness. By contrast, increasing shock durations produced greater conditioned fear to the shock context than did decreasing shock durations in Experiment 2. The differential effects of shock-duration order on finickiness and fear are explicated in terms of the specificity of fear conditioning during exposure to increasing versus decreasing series of shock duration orders.     

Influence of conspecific stress odors and shock controllability on conditioned defensive burying

In Experiment 1, rats received a session of 80 inescapable tail shocks or no shocks while restrained in a tube. During tests of conditioned defensive burying 24 h later, the bedding of the chamber contained odors from either stressed or nonstressed conspecific donor rats. Following a single prod shock, subjects that had had prior shocks or that were tested with the stress odors spent significantly less time burying the prod, made smaller piles of bedding, and displayed more freezing behavior. The combination of prior shock and stress odors during later testing enhanced these effects. In Experiment 2, a yoked group of rats that was given inescapable shocks, in contrast to a group that had wheel-turn escape training and one that was restrained but not shocked, later showed significantly less burying and more freezing when tested for defensive burying with stress odors present. In both experiments the duration of burying and the heights of piles were positively correlated, and both of these measures were negatively correlated with freezing. The demonstrated capacity of unconditioned stress odors to mediate different degrees of fear, depending upon the controllability of prior shock, is related to other studies of learned helplessness, and the predominance of freezing over burying is discussed in terms of various types of defensive strategies, stimulus-control processes, and the author’s stress-coping-fear-defense (SCFD) theory.     

Control over noxious events and choice

Three experiments are reported assessing whether rats prefer controllable over uncontrollable aversive shock. In Experiment 1, subjects chose between escapable and inescapable shock while relative shock duration varied parametrically. In Experiment 2, subjects again chose between escapable and inescapable shock, but duration was held constant and equal. The final experiment gave subjects a choice between avoidable and unavoidable shock under several signaling conditions. Choice behavior proved sensitive to relative shock duration and to predictability of shock but not to controllability of shock.     

Preexposure to situational cues and shock intensity in two-way avoidance learning

Four groups of rats (n = 16) received 65 two-way avoidance learning trials. The groups differed with respect to the amount of exposure (0 or 4 h) to the situational cues of the apparatus prior to avoidance learning and the intensity of shock (.3 or 1.6 mA) during learning. Superior avoidance performance with weak as compared to strong shock was obtained in the nonpreexposed groups. This inverse relationship between avoidance performance and shock intensity, typical of two-way avoidance learning, was eliminated in the preexposed groups. Presumably, a latent inhibition effect occurred in the strong-shock group, which resulted in a retardation of the conditioning of fear to the situational cues and a consequent improvement in performance. The results are consistent with the effective reinforcement theory, which emphasizes in aversive learning the detrimental effect of large amounts of fear remaining following a response.     

Preexposure to situational cues produces a direct relationship between two-way avoidance learning and shock intensity

In Experiment 1, four groups of subjects (n = 16 each) were exposed to the situational stimuli of a shuttlebox apparatus for 4 h. Subsequently, 200 two-way avoidance trials were administered (100/day) with either .3- or 1.6-mA shock and with either small or large reward (presence or absence of visual stimuli following the response). Avoidance performance was directly related to shock intensity on both days and to magnitude of reward on the 2nd day. In Experiment 2, four groups of subjects (n = 24 each) were given 4 h of exposure either to the situational stimuli of the shuttlebox or to a neutral box. Then, 10 two-way avoidance trials were given with 1.6-mA shock. Subsequently, subjects were allowed to escape from one of the shuttlebox compartments to an adjacent safe box. Following preexposure to situational stimuli, avoidance performance was superior whereas escape-from-fear performance was inferior. This latter finding demonstrated that less fear of situational cues was present during avoidance training in the preexposed condition. All of these results support the effective reinforcement theory, an extension of two-factor theory, which emphasizes the importance for avoidance learning of the amount of fear of situational cues present following a response.     

Effects of cuing in an “irrelevant” context

Normally, retention of an avoidance response by a rat is impaired when the test context is novel or does not correspond to the training context. Experiment 1 demonstrates that such an impairment of test performance can be alleviated if a rat receives a cuing treatment or reminder of training in the novel test context prior to testing. Experiment 2 indicates that when rats receive avoidance training in one context and then receive a reminder of training in a novel context, they perform more poorly when tested in the training context than do animals that receive no reminder. This finding is discussed in relation to current theories of contextual influence over retention performance.     

Shock vs. ice-water passive avoidance learning in wild and domesticRattus norvegicus

Two experiments comparing passive avoidance responses of wild and domestic Norway rats included both shock and ice water as aversive stimuli. Experiment I indicated that wild rats did not exhibit passive avoidance in these situations and that shock was more effective than ice water as an aversive stimulus for domestic rats. Modifications of the ice-water apparatus in Experiment II resulted in improved passive responses in domestic rats, but ice water was still not as effective as shock. Wild rats again showed no passive avoidance. The results of these two experiments were interpreted in terms of stock differences in species-specific defense reactions for these avoidance situations.     

Reinstatement of fear to an extinguished conditioned context

Rats were shocked in the black but not the white compartment of a shuttlebox and then exposed to the black compartment in the absence of the shock unconditioned stimulus (US) to extinguish fear responses (passive avoidance). In five experiments, rats were then shocked in a reinstatement context (distinctively different from the shuttlebox) to determine the conditions that reinstate extinguished fear responding to the black compartment. Rats shocked immediately upon exposure to the reinstatement chamber failed to show either reinstatement of avoidance of the black compartment or fear responses (freezing) when tested in the reinstatement chamber. In contrast, rats shocked 30 sec after exposure to the reinstatement chamber exhibited both reinstatement of avoidance of the black compartment and freezing responses in the reinstatement chamber (Experiment 1). Rats shocked after 30 sec of exposure to the reinstatement chamber but then exposed to that chamber in the absence of shock failed to exhibit reinstatement of the avoidance response and did not freeze when tested in the reinstatement chamber (Experiment 2). Rats exposed to a signaled shock in the reinstatement chamber and then exposed to that chamber in the absence of shock also failed to exhibit reinstatement of the avoidance response (Experiment 5). These rats showed fear responses to the signal but not to the reinstatement chamber. Finally, rats exposed for some time (20 min) to the reinstatement chamber before shock exhibited reinstatement of the avoidance response but failed to freeze when tested in the reinstatement chamber (Experiments 3 and 4). These results are discussed in terms of the contextual conditioning (Bouton, 1994) and the US representation (Rescorla, 1979) accounts of postextinction reinstatement.     

Effects of stress controllability,immunization, and therapy on the subsequent defeat of colony intruders

Three experiments investigated the influence that various stress-controllability manipulations had on the defensive behaviors of rats when they were subsequently tested as intruders in previously established, aggressive colonies of conspecifics. In Experiment 1, naive subjects that had received a session of 80 shocks in a tube showed an enhanced series of defensive responses and received more bites than did a group of restrained nonshocked rats as colony intruders 24 h later. These two measures were also found to be positively correlated within each group. In Experiment 2, a group that was given 80 yoked inescapable shocks, in contrast to a group that had wheel-turn escape training and a restrained nonshocked control group, displayed more defeat and was bitten more frequently when tested as intruders on the following day. In Experiment 3, 60 trials of wheel-turn escape training were given 4 h prior to (i.e., immunization) or after (i.e., therapy) a session of 60 inescapable tube shocks. During resident-intruder testing 24 h later, both of these groups showed less defeat and received fewer bites than did an inescapably preshocked group but did not differ from a restrained nonshocked control group. These findings clearly indicate that stress controllability alters species-typical defensive responses, and their implications concerning other learned helplessness effects and interpretations are discussed.     

Continuity and intensity of shock in one-way avoidance learning in the rat

Six groups of rats (n = 16) differed with respect to the continuity of shock (continuous or discontinuous) and the shock intensity (.3, .8, or 1.6 mA) used during 65 one-way avoidance-conditioning trials. In general, a facilitative effect on one-way avoidance learning was obtained for continuous as opposed to discontinuous shock and for strong as opposed to weak shock. For both variables, the results are opposite to those obtained in discriminated shuttlebox-avoidance and barpress avoidance tasks. The data support an interpretation of the effect of continuity of shock which holds that discontinuous shock is, in effect, less intense than continuous shock. This interpretation allows the effects of the continuity-of-shock variable to be incorporated within the effective reinforcement theory of avoidance learning which has been proposed to account for shock-intensity effects in various avoidance tasks.     

Discriminated and nondiscriminated avoidance conditioning of the rearing response in rats

Experiment 1 employed a shock box in which light beams ran at 10, 15, 20, or 25 cm above the floor level of the box. Four groups of nine rats each were trained to avoid shock by cutting the light beams or letting them pass by, which the animal accomplished by upward or downward change of its posture. Training employed a discriminated avoidance paradigm, 60 trials per day for 5 days, with a 5-sec CS-US interval. Acquisition of the rearing avoidance response was observed only in the 15-cm condition. Using the same apparatus as in Experiment 1 and with a beam height of 15 cm, the rearing avoidance response was successfully conditioned in five rats using a nondiscriminated avoidance conditioning paradigm. There was good evidence of temporal discrimination in these animals.