Developments of a water-maze procedure for studying spatial learning in the rat

\, SUMMARY In 1967, Overmier and Seligman found that dogs exposed to inescapable and unavoidable electric shocks in one situation later failed to learn to escape shock in a different situation where escape was possible. Shortly thereafter Seligman and Maier (1967) demonstrated that this effect was caused by the uncontrollability of the original shocks. In this article we review the effects of exposing organisms to aversive events which they cannot control, and we review the explanations which have been offered. There seem to be motivational, cognitive, and emotional effects of uncontrollability. (a) Motivation. Dogs that have been exposed to inescapable shocks do not subsequently initiate escape response in the presence of shock. We review parallel phenomena in cats, fish, rats, and man. Of particular interest is the discussion of learned helplessness in rats and man. Rats are of interest because learned helplessness has been difficult to demonstrate in rats. However, we show that inescapably shocked rats do fail to learn to escape if the escape task is reasonably difficult. With regard to man, we review a variety of studies using inescapable noise and unsolvable problems as agents which produce learned helplessness effects on both instrumental and cognitive tasks, (b) Cognition. We argue that exposure to uncontrollabl e events interferes with the organism's tendency to perceive contingent relationships between its behavior and outcomes. Here we review a variety of studies showing such a cognitive set. (c) Emotion. We review a variety of experiments which show that uncontrollable aversive events produce greater emotional disruption than do controllable aversive events. We have proposed an explanation for these effects, which we call the learned helplessness hypothesis. It argues that when events are uncontrollable the organism learns that its behavior and outcomes are independent, and that this learning produces the motivational, cognitive, and emotional effects of uncontrollabi lity. We describe the learned helplessness hypothesis and research which supports it. Finally, we describe and discuss in detail alternative hypotheses which have been offered as accounts of the learned helplessness effect. One set of hypotheses argues that organisms learn motor responses during exposure to uncontrollabl e shock that compete with the response required in the test task. Another explanation holds that uncontrollable shock is a severe stressor and depletes a neurochemical necessary for the mediation of movement. We examine the logical structure of these explanations and present a variety of evidence which bears on them directly.

Learned helplessness: Theory and evidence.

We examine two different descriptions of the behavioral functions of the hippocampal system. One emphasizes spatially organized behaviors, especially those using cognitive maps. The other emphasizes memory, particularly working memory, a short-term memory that requires iexible stimulus-response associations and is highly susceptible to interference. The predictive value of the spatial and memory descriptions were evaluated by testing rats with damage to the hippocampal system in a series of experiments, independently manipulating the spatial and memory characteristics of a behavioral task. No dissociations were found when the spatial characteristics of the stimuli to be remembered were changed; lesions produced a similar deficit in both spatial and nonspatial test procedures, indicating that the hippocampus was similarly involved regardless of the spatial nature of the task. In contrast, a marked dissociation was found when the memory requirements were altered. Rats with lesions were able to perform accurately in tasks that could be solved exclusively on the basis of reference memory. They performed at chance levels and showed no signs of recovery even with extensive postoperative training in tasks that required working memory. In one experiment all the characteristics of the reference memory and working memory procedures were identical except the type of memory required. Consequently, the behavioral dissociation cannot be explained by differences in attention, motivation, response inhibition, or the type of stimuli to be remembered. As a result of these experiments we propose that the hippocampus is selectively involved in behaviors that require working memory, irrespective of the type of material (spatial or nonspatial) that is to be processed by that memory.

https://www.cambridge.org/core/services/aop-cambridge-core/content/view/F9ABFA09F5AC96A64D15572521BD2C6C/S0140525X00062713a.pdf/div-class-title-hippocampus-space-and-memory-div.pdf

Hippocampus, space, and memory

The history of 2-process learning theory is described, and the logical and empirical validity of its major postulates is examined. The assumption of 2 acquisition processes requires the demonstration of an empirical interaction between 2 types of reinforcement contingencies and (a) response classes, (b) reinforcing stimulus classes, or (c) characteristics of the learned behavior itself. The mediation postulates of 2-process theory which argue that CRs are intimately involved in the control of instrumental responding are emphasized, and 2 major lines of evidence that stem uniquely from these postulates are examined : (a) the concurrent development and maintenance of instrumental responses and conditioned reflexes, and (b) the interaction between separately conducted Pavlovian conditioning contingencies and instrumental training contingencies in the control of instrumental behavior. The evidence from concurrent measurement studies provides, at the very best, only weak support for the mediational hypotheses of 2-process theory. In contrast, the evidence from interaction studies shows the strong mediating control of instrumental responses by Pavlovian conditioning procedures, and demonstrates the surprising power of Pavlovian concepts in predicting the outcomes of many kinds of interaction experiments.

Two-process learning theory: Relationships between Pavlovian conditioning and instrumental learning

The present experiments were designed to examine the hypothesis that the mammalian brain contains anatomically distinct memory systems. Rats with bilateral lesions of caudate nucleus or fimbria-fornix and a control group were tested postoperatively on 1 of 2 versions of the radial maze task. In a standard win-shift version, each of the 8 arms of the maze was baited once, and the number of errors (revisits) in the first 8 choices of each trial was recorded. Fimbria-fornix rats were impaired in choice accuracy, while caudate animals were unimpaired relative to controls. Different groups of rats with similar lesions were tested on a newly developed win-stay version of the radial maze, in which the location of 4 randomly selected baited arms was signaled by a light at the entrance to each arm, and which required rats to revisit arms in which reinforcement had been previously acquired within a trial. Rats with fimbria-fornix lesions were superior to controls in choice accuracy on the win-stay radial maze task, while caudate animals were impaired relative to controls. The results demonstrate a double dissociation of the mnemonic functions of the hippocampus and caudate nucleus. Some implications of the presence of 2 memory systems in the mammalian brain are discussed.

https://www.jneurosci.org/content/jneuro/9/5/1465.full.pdf

Differential effects of fornix and caudate nucleus lesions on two radial maze tasks: evidence for multiple memory systems.

Stimulus control of spatial behavior on the eight-arm maze in rats ☆ ☆☆

The effects of restraint and noncontingent preshock on subsequent escape learning in the rat

Heart rate of rats was recorded in the Estes-Skinner conditioned emotional response situation. Response to the conditioned stimulus was a decrease in rate. The change in heart rate was conditioned more slowly than suppression of bar-pressing; it was of shorter duration and was more variable than suppression.

https://science.sciencemag.org/content/sci/152/3727/1404.full.pdf

Heart Rate: Changes during Conditioned Suppression in Rats

Dogs were trained to press a pedal to avoid shock. They were then operantly reinforced for making or for refraining from making a series of electromyographic responses while almost completely curarized. Tests after recovery from curarization showed that operant conditioning under curare influenced the original pedal-press response.

http://science.sciencemag.org/content/sci/155/3759/201.full.pdf

Transfer Following Operant Conditioning in the Curarized Dog

Retention of aversively motivated responses in rats.

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