https://numerons.files.wordpress.com/2012/04/11-stressor-controllability-and-learned-helplessness.pdf

Stressor controllability and learned helplessness: the roles of the dorsal raphe nucleus, serotonin, and corticotropin-releasing factor.

Learned helplessness, the failure to escape shock induced by uncontrollable aversive events, was discovered half a century ago. Seligman and Maier (1967) theorized that animals learned that outcomes were independent of their responses-that nothing they did mattered-and that this learning undermined trying to escape. The mechanism of learned helplessness is now very well-charted biologically, and the original theory got it backward. Passivity in response to shock is not learned. It is the default, unlearned response to prolonged aversive events and it is mediated by the serotonergic activity of the dorsal raphe nucleus, which in turn inhibits escape. This passivity can be overcome by learning control, with the activity of the medial prefrontal cortex, which subserves the detection of control leading to the automatic inhibition of the dorsal raphe nucleus. So animals learn that they can control aversive events, but the passive failure to learn to escape is an unlearned reaction to prolonged aversive stimulation. In addition, alterations of the ventromedial prefrontal cortex-dorsal raphe pathway can come to subserve the expectation of control. We speculate that default passivity and the compensating detection and expectation of control may have substantial implications for how to treat depression. (PsycINFO Database Record

https://psycnet.apa.org/journals/rev/123/4/349.pdf

Learned helplessness at fifty: Insights from neuroscience.

Animal models of depression: an overview.

Some animal models of depression, including the majority of the more recently introduced models, are better characterized as models of predisposition to depression. In the first part of this paper, we show that the basis for such a model could be either a procedure that increases the ease with which an analogue of major depression may be evoked, or a presentation analogous to dysthymia (chronic mild depression). We then consider how the concepts of predictive, face, and construct validity apply to such models. Next, we review the validity of the available models of predisposition to depression, which derive from genetics, genomics, developmental manipulations, and brain lesioning. Finally, we compare the performance of the different models, using a novel scoring system that formalizes the evaluation of animal models against each of the three sets of validation criteria.

The validity of animal models of predisposition to depression.

The degree to which organisms can exert control over events to which they are exposed has a strong impact on behavior and physiological functioning. Effects caused by the uncontrollability of events that are beyond the organism's control rather than by the events per se have been called learned helplessness effects. The present paper reviews such learned helplessness effects. At a behavioral level, uncontrollable aversive events result in associative, motivational, and emotional deficits. At a neurochemical level, uncontrollable but not controllable aversive events have been reported to lead to disturbances in cholinergic, noradrenergic, dopaminergic, serotonergic, and GABAergic systems. However, there are interpretive difficulties in this literature, and these are discussed. The controllability/uncontrollability of aversive events has a role in producing stress-induced analgesia and the activation of endogenous opiate systems. These relationships are reviewed. It is proposed that the learning that aversive events cannot be controlled activates an opiate system. The research reviewed is related to depression, and the general issue of animal models of depression is discussed. It is concluded that no experimental paradigm can be a model of depression in some general sense, but can only model a particular aspect. Learned helplessness may model "stress and coping".

Learned helplessness and animal models of depression.

Exposure to inescapable shock produces both activity and associative deficits in the rat

The role of temporal factors in the development of conditioned inhibition was investigated in a backward conditioning design. Separate groups of rats received tone CSs either 3 or 30 sec following shock presentatlOns. The ess predicted the same shock-free interval for both groups. A third group was presented with a random relationship between CS and shock. The CSs were tested by super­ imposition on a Sidman avoidance baseline and only the group with a 3-sec UCS·CS interval revealed an inhibitory effect of the CS. These results are in accord with predictions made by the Solomon-Corbit model of acquired motivation and by Denny's "relaxation" theory of escape and avoidance.

/pdf/conditioned-inhibition-and-the-ucs-cs-interval-1etulpomzm.pdf

Conditioned inhibition and the UCS-CS interval.

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.

/pdf/inescapable-shock-and-food-competition-dominance-in-rats-4ru32ke5in.pdf

Inescapable shock and food-competition dominance in rats

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Exposure to inescapable shock produces both activity and associative deficits in the rat

Conditioned inhibition and the UCS-CS interval.

Inescapable shock and food-competition dominance in rats