seligman's learned helplessness experiment

Kerr DL, McLaren DG, Mathy RM, Nitschke JB. The non-difference between the zero group and the escapable shock group led us to believe that organisms expected controllability as the basic default option. Peyron C, Petit JM, Rampon C, Jouvet M, Luppi PH. Ray RD, Zald DH. Hiroto DS, Seligman MEP. For complex organisms behavioral control can be possible over threats that are repeated, intermittent, and so persist across time. Although these descending neurons are glutamatergic and so excitatory, they synapse preferentially on GABAergic interneurons in the dorsal raphe nucleus that inhibit the 5-HT cells (see Figure 2 for a cartoon). This suggests that if the DETECT control circuit is absent the animal invariably reverts to the default of helplessness following exposure to any prolonged stressor. Perceived controllability modulates the neural response to pain. Retrograde and anterograde tracing studies indicate that the dorsal raphe nucleus does not send a major projection to the paraventricular nucleus of the hypothalamus, and perhaps none at all (Larsen, Hay-Schmidt, Vrang, & Mikkelsen, 1996). One might ask if medial prefrontal cortical activity inhibits the dorsal raphe nucleus when therapy or medication is successful. To find out they restrained dogs in a hammock and the dogs got 64 mild-moderate electric shocks to their back paws, each shock heralded by a tone. It is a circuit that provides an expectational function, in the sense that it changes or biases how organism's respond in the future as a consequence of the events that occur in the present. But, if control leads to EXPECT (sensitized prelimbic-dorsal raphe nucleus inhibition), why should the escapably shocked subjects not perform better than controls that had not been previously stressed and so do not have EXPECT? So it is important to distinguish between what the brain does under a given set of conditions, and what the brain is capable of under different conditions. Celebrate? Maier SF, Ryan SM, Barksdale CM, Kalin NH. Adrenocorticotrophic hormone in turn stimulates the production and release of glucocorticoids (corticosterone in the rat, cortisol in humans) from the adrenal cortex into the blood. The dorsal raphe nucleus sends 5-HT projecting neurons to numerous regions including the periaqueductal gray, striatum, and extended amygdala. Third, is activation of this pathway sufficient for control to reduce dorsal raphe nucleus activation and block the passivity typically produced by inescapable shock? Grahn RE, Watkins LR, Maier SF. Anatomical insights into the interaction of emotion and cognition in the prefrontal cortex. However, these reflexes are energy intensive, and so if unsuccessful it might be adaptive to inhibit them and conserve energy for use in physiological adjustments that promote survival, such as altering the responsivity of the immune system to be better able to fight any infection or wound that might occur after an attack (Frank, Watkins, & Maier, 2013). They also planned ways to prevent self-harm, and they planned the ways that he would capitalize on his progress in the coming year and they explored how he could continue to be in a good place one year from now. When common cues are shared between the first environment and the test environment, processes such as fear conditioning could mediate the behavioral change. As support, the addition of external stimuli such as a light or a tone occurring immediately after each INESC prevented the occurrence of later failure to learn to escape (Minor et al., 1990; Weiss, 1971). Importantly, the presence of control aborts this process. Seligman MEP. Of course, inescapable shock produces a greater activation of the dorsal raphe nucleus than does controllable shock. In addition to theory testing, knowledge of the underlying circuitry explained a number of learned helplessness phenomena that were simply mysteries at a psychological level. Although there is a long history of research investigating the controllability dimension in humans, studies using methods that allow the measurement of neural activity are quite recent and few in number. The presence of control could reduce or inhibit paraventricular activation only if the structures that DETECT control project to the paraventricular nucleus. To do so the dorsal raphe nucleus would have to extract the conditional probability of the shock offset given that the wheel turn or some other escape response occurs, and the conditional probability of the shock offset occurring in the absence of those responses, and compare these two probabilities. Cognitive therapy versus medication for depression: Treatment outcomes and neural mechanisms. Martin Seligman and Steven F. Maier first identified learned helplessness as a phenomenon in the 1960s. Activation of serotonin-immunoreactive cells in the dorsal raphe nucleus in rats exposed to an uncontrollable stressor. Here are the mechanisms that were assumed by the original theory: First: DETECT: Animals DETECT the dimension of controllability and uncontrollability. The first blush reaction is that we should measure these structures in humans, and then excite the ventromedial prefrontal cortex and inhibit the dorsal raphe nucleus, pharmacologically, electrically, trans-magnetically or psychologically in therapy. What do you do? That is, the findings that we have reviewed do not imply that the brain does not have circuitry to detect non-contingency between events that include actions and outcomes. In the only relevant triadic design of which we are aware, Kerr, McLaren, Mathy, and Nitschke (2012) used exposure to snake videos to subjects with snake phobias. Figure 1 shows the levels of 5-HT within the dorsal raphe nucleus during escapable and inescapable shock. The next section provides a brief summary of this early work. However, the speed required on controllable trials was adjusted so that the subjects succeeded about half the time in avoiding the video, and so the actual frequencies on the uncontrollable trials was equated to this frequency. The role of the amygdala in fear and conditioned fear is well known. This work led us to define a dimension that we called control over outcomes, with control being present whenever the probability of (O), given a response (R) is different from the probability of (O) given the absence of that response (notR). But the neural data are clear. Rozeske RR, Evans AK, Frank MG, Watkins LR, Lowry CA, Maier SF. Control over a stressor involves the posterior dorsal striatum and the act/outcome circuit. 8-OH-DPAT microinjected in the region of the dorsal raphe nucleus blocks and reverses the enhancement of fear conditioning and the interference with escape produced by exposure to inescapable shock. Here, an expectancy of control does blunt the impact of subsequent stressors. Attentional control and mindfulness similarly emphasize the present. The site is secure. Journal of Comparative Physiology A: Neuroethology, Sensory, Neural, and Behavioral Physiology. All this suggested that perhaps DETECTion of control over shock is done by a circuit involving the prelimbic area and the dorsal medial striatum, just as is instrumental appetitive contingency learning. Donald S. Hiroto and Martin E. P. Seligman University of Pennsylvania Learned helplessness, the interference with instrumental responding followinginescapable aversive events, has been found in animals and man. With knowledge of the underlying neural circuitry it became simple to ask whether safety signals blunt the impact of stressors via the same or a different mechanism. Moreover, neurons in the ventromedial prefrontal cortex participate in numerous circuits with neurons in other brain regions, and the functions served by these circuits are likely unrelated, except that they share cells in a large piece of heterogeneous geography. This is a cumulative process with 5-HT building over time, and if the transmitter reaches some threshold in target structures, defensive behaviors become inhibited and energy flow is shifted. Title. He was afraid he would be distraught on this anniversary and engage in self-harm. Shin LM, Liberzon I. Seligman MEP, Csikszentmihalyi M. Positive psychology: An introduction. Thus, subjects were exposed to escapable shock or yoked inescapable shock in the wheel-turn apparatus and given fear conditioning in standard conditioning chambers seven days later. The cells in this region are GABAergic and project to and inhibit the central nucleus of the amygdala (Berretta, Pantazopoulos, Caldera, Pantazopoulos, & Pare, 2005). The technique of in vivo microdialysis allows the measurement of the levels of 5-HT in discrete brain regions in real-time in live, awake, behaving animals. Reappraisal as a tool of therapy is behavioral control. (2006) noted that these regions of the prefrontal cortex do not project to the amygdala, but they do project to the ventromedial prefrontal cortex. Importantly, control still blunts the impact of the stressor being experienced, but longer-term immunization is eliminated; 2) The production of particular proteins, called plasticity proteins, is required for long term-increases in connectivity. In this paper for simplicity, we will refer to this suite of changes as passivity/anxiety, but we emphasize that we are interested in all of the changes separatelybut not for the purpose of the present theorizing. However, only roughly 1/3 of dorsal raphe nucleus neurons contain 5-HT, and so simply showing generalized activation is not enough. This was exactly why we used the triadic design and included the escapable subjects as a control group because it isolated the element of uncontrollabilityif failure to escape in the shuttlebox was caused by learning uncontrollability, then this failure should not occur if uncontrollability is removed but shock stays constant. Blockade of 5-HT receptors in the dorsal raphe nucleus target regions during later testing. To answer this question M. Barratta (unpublished) in Maier's group microinjected one color retrograde tracer in the dorsal medial striatum and a differently colored retrograde tracer into the dorsal raphe nucleus. This is likely because running is elicited as a species specific defense response (Bolles & Fanselow, 1980) on the very first trials. However, this does not mean that control is necessarily detected by the ventromedial prefrontal cortex in isolation, or by the ventromedial prefrontal cortex at all. Recent evidence supports this idea: 1) The development of persistent increases in connectivity requires the production of new proteins in the cells in question, and blockade of new protein synthesis in the prelimbic area after the escapable shock experience prevents immunization (Amat et al., 2006). Radley JJ, Sawchenko PE. When this is true of all responses, objective helplessness exists. These project to the dorsal raphe nucleus, and consistent with the Weiss work, blockade of NEreceptors in the dorsal raphe nucleus with a microinjected antagonist during inescapable shock eliminated the passivity and anxiety (Grahn et al., 2002). Primitive organisms do not have the sensory apparatus to detect threats at a distance, nor do they have a complex behavioral repertoire that can be used for what we are calling behavioral control. Chlordiazepoxide microinjected in the region of the dorsal raphe nucleus eliminates the interference with escape responding produced by inescapable shock whether administered before inescapable shock or escape testing. Failure to escape traumatic shock. Well, she replied, you've obviously come a really long way since then: You're working again, you're in a great relationship. The psychology of depression: Contemporary theory and research. Martin E. P. Seligman, University of Pennsylvania. Here are two examples. Note that by this view there is no learning about uncontrollability or controllability, just Pavlovian processes and fear. Recall that the phenomenon that began this line of work was that exposure to aversive Pavlovian fear conditioning leads to later failure to learn instrumental escape/avoidance responses (Leaf, 1964). Baratta et al. The two-factor theory of avoidance learning predicted that turning on the fear-inducing tone in the shuttlebox would generate fear and accelerate jumping. (2010) reported that experience with escapable shock blocked the passivity in shuttle escape and also blocked the reduced social investigation (anxiety) produced by social defeat occurring seven days later. This suggests that the rat EXPECTs that shock will be escapable in the new situation and that plasticity in the prelimbic-dorsal raphe nucleus subserves this expectation and inhibits the dorsal raphe nucleus, thus blocking learned helplessness. Quiz & Worksheet Goals. We arrived at Penn as first year graduate students in 1964. First, baseline levels were measured before the stressors began. A number of studies employing painful stimuli have found that providing control, or inducing perceived control, reduces the experienced intensity of the painful stimulus. Avoidance response evocation as a function of prior discriminative fear conditioning under curare. government site. Prefrontal control of resilience to adverse events. The results were dramatic. Seligmans Learned Helplessness Theory, Explained. To test this idea Amat et al. That is, if the control detecting circuit was taken off line, all animals acted as if the shock was inescapable whether the animal actually was able to escape the shock or not. Control could be detected by a different circuit, with this information then conveyed over to the ventromedial prefrontal cortex. However, the mere presence of a juvenile leads to a large increase in 5-HT within the amygdala in a subject that has experienced inescapable shock, but not escapable shock previously (Christianson et al., 2010). We thought that a profound failure to escape was the phenomenon and we began to try to understand it. Dramatically, inescapable shock now does activate these cells as if the inescapable shock was escapable shock if the organism had experienced immunizing control seven days earlier! Special thanks go to J. Amat, S. Bland, M. Baratta, J. Christianson, A. Der-Avakian, R. Drugan, J. Elstein, R. Grahn, J. Hammack, R.L. Differential projections of the infralimbic and prelimbic cortex in the rat. Instead, she played dumb: Wow, the one-year anniversary. If control leads to sensitization of the prelimbic-dorsal raphe nucleus pathway (EXPECT), then 5-HT activity should be inhibited from the start during shuttlebox escape testing in this group, but not in the non-shocked controls. https:// https://doi.org/10.1037/0096-3445.105.1.3 Abstract Reviews the literature which examined the effects of exposing organisms to aversive events which they cannot control. The most proximate mediator of fight/flight seemed to be the dorsal periaqueductal gray (dPAG), while the extended amygdala (bed nucleus of the stria terminalis, BNST, together with the amygdala proper) mediated fear/anxiety. In sum, a number of inputs to the dorsal raphe nucleus, using a number of different transmitters, was necessary to produce learned helplessness behaviors, but these inputs did not discriminate inescapable from escapable shock. As this conclusion also predicts, inactivating the dorsolateral striatum and the habit system did not reduce the protective effects of escapability. This circuit is about the future and it buffers against, but does not annihilate, the default reaction of the dorsal raphe nucleus. Role of the ventral medial prefrontal cortex in mediating behavioral control-induced reduction of later conditioned fear. Importantly, behavioral sequelae of IS such as escape deficits and anxiety also persist for just this period of time (Maier, 2001). The detection of uncontrollability is not necessary nor is it sufficient for passivity. This requires a method that allows the experimenter to know that a prelimbic neuron that is being recorded is in one of these two pathways (most prelimbic neurons are in neither), and which one. There was no colocalization at all, indicating that these are separate populations of prelimbic neurons. That is, even though the subjects exert control and the prelimbic area is activated, immunization only happens if new proteins can be formed. Goldin PR, Ziv M, Jazaieri H, Hahn K, Heimberg R, Gross JJ. There is a growing and complex literature concerning the impact of therapies such as cognitive-behavioral therapy (CBT) on neural function that cannot be reviewed here. Instead, safety signals had their impact via the insular cortex and insular cortex lesions eliminated the protective effects of safety signals. Assuming that learned helplessness models these phenomena, we now speculate on the implications of the neural circuitry particularly for the treatment of depression. Whether the ventromedial prefrontal cortex activation produced by reappraisal involves the specific pathways that are critical to mediating the impact of behavioral control awaits future research. FOIA Reversal of learned helplessness by imipramine. However, both escapable and inescapable shock produced exactly equal levels of locus coeruleus NE activation (McDevitt et al., 2009). However, it should be clearly understood that this EXPECTATION may not be a cognitive process or entity as psychologists tend to view them. Third: EXPECT. At the psychological level, there are several other loose ends. Annual Review of Medicine Vol. The video or a fixation point then appeared. Activation of this structure might then recapitulate the behavioral pattern produced by inescapable shock. In a nutshell learned helplessness is when a person (or animal) faces negative, uncontrollable circumstances and stops trying to improve upon their situation even when they can. This is accomplished by a circuit involving projections from the prelimbic region of the ventromedial prefrontal cortex (the PL) to the dorsal medial striatum (the DMS), and back. Maren S, Quirk GJ. As a general statement, neural processes in the prefrontal cortex become narrowed by stress (Arnsten, 2015). She found that when subjects reduced negative emotional reactions successfully, there was a strong negative correlation between amygdala and ventromedial prefrontal cortex activity. In the rodent, the corticosterone rise is not greater or more prolonged (Helmreich et al., 2012; Maier et al., 1986), the adrenocorticotrophic hormone rise is not greater or more prolonged (Maier et al., 1986), nor is the increase in corticotropin releasing hormone in the hypothalamus larger (Helmreich et al., 1999). The reader may wonder why then in all the initial helplessness experiments the previously nonshocked group and the previously escapably shocked group performed equally well in shuttlebox escape. We now go through the neural circuitry dataset in detail using the PASSIVITY/ANXIETY, DETECT, ACT, and EXPECT terminology both to make the argument more easily understood and because we believe that it is a useful translation from the neural level of analysis to the psychological level of analysis. There are numerous consequences of exposure to an aversive event that are, in fact, not modulated by control (Helmreich et al., 1999; Maier, Ryan, Barksdale, & Kalin, 1986; McDevitt, et al., 2009; Woodmansee, Silbert, & Maier, 1993). There is no question that the presence of safety signals that predict a period of time free from shock can reduce the behavioral impact of aversive events. One system, called the act/outcome system, is sensitive to the contingency between response and reinforcer. Maier SF, Grahn RE, Watkins LR. This expectation undermined their trying (response initiation) to escape. Research with animals quickly switched from dogs to rodents, but using the same triadic design that compares exactly equal ESC, yoked INESC, and no shock. Amat J, Paul E, Zarza C, Watkins LR, Maier SF. Greenwood BN, Strong PV, Fleshner M. Lesions of the basolateral amygdala reverse the long-lasting interference with shuttle box escape produced by uncontrollable stress. Inescapable shock now activates the sensitized prelimbic- dorsal raphe nucleus pathway, which now operates as an EXPECT circuit. Effect of inescapable shock on subsequent escape performance: Catecholaminergic and cholinergic mediation of response initiation and maintenance. Prospection: Experiencing the future. Sierra-Mercado D, Padilla-Coreano N, Quirk GJ. When shock is initially escapable, the presence of control is DETECTed. However, it was never entirely clear how or why increased activity of these neurons would produce the passivity or anxiety. (2005) inactivated the ventromedial prefrontal cortex-to-dorsal raphe nucleus pathway during the experience of escapable shock, inescapable shock, or no shock. Does the ventromedial prefrontal cortex actually regulate dorsal raphe nucleus activity and passivity as specified by this model (Figure 3)? This is now a circuit that EXPECTS control. In these studies, one group (1) of dogs was given repeated, inescapable shocks while restrained. Indeed, Greenwood, Strong, and Fleshner (2010) have shown this to be the case. and transmitted securely. Clearly, if such control will work this is the best course because it will minimize injury and harm. i10-index. As organisms became more complex, they could detect and identify threats at a distance. Learned stressor resistance requires extracellular signal-regulated kinase in the prefrontal cortex. However, the work of J. Weiss was an exception. Thus, DETECT escapability and ACT to pass this information that then inhibits the dorsal raphe nucleus are subserved by different populations of ventromedial prefrontal cortex cells and so are truly different functions. With regard to sufficiency, simply activating the dorsal raphe nucleus by microinjecting agents into the dorsal raphe nucleus that activate 5-HT neurons should produce the same passivity and anxiety as does inescapable shock. First: PASSIVITY/ANXIETY. So in the original stressor controllability experiment (Seligman & Maier, 1967), this group of dogs learned in the hammock to press a panel with their noses to turn off each shock. This assertion could be tested, although the appropriate experiment has not yet been done. Changing fear: The neurocircuitry of emotion regulation. Activation of the ventral medial prefrontal cortex during an uncontrollable stressor reproduces both the immediate and long-term protective effects of behavioral control. However, if control is possible this is detected and leads to the inhibition of this process so that active responding can continue. This experiment was one of several studies throughout the 1970s, 1980s, and 1990s that laid the foundation for a new theory of human behavior related to failure, learned helplessness, and resilience. We conclude from this that the prelimbic-dorsal raphe nucleus ACT pathway can be modified over several hours after the joint experience of control and aversive stimulation, to respond to stressors in general as if they were controllable, and this is compatible with the idea that this altered pathway subserves the EXPECTation that shock will be controllable in new aversive situations. Balleine BW, Dickinson A. Goal-directed instrumental action: Contingency and incentive learning and their cortical substrates. If the behavioral effects of inescapable shock are mediated by the learned expectation that active responding will not produce relief, the original idea, then why this time course? During later inescapable shock, the ESC group which had first learned to bar press to escape shock continued to run in a shuttlebox during long duration inescapable shock. Helmreich DL, Tylee D, Christianson JP, Kubala KH, Govindarajan ST, O'Neill WE, Maier SF. The neural evidence strongly suggests that activation and sensitization of the dorsal raphe nucleus leads to the passivity and anxiety characteristic of learned helplessness. Neuroscience research indicates that each can take place and each involves different neural systems (Balleine & O'Doherty, 2010). Why should control modulate passivity, fear, and the neurochemical impact of a stressor but not the hypothalamo-pituitary-adrenal response? Citation Maier, S. F., & Seligman, M. E. (1976). Monamines as mediators of avoidance-escape behavior*. Second, the initial stressor must occur in an environment very different from the test environment since one of the hallmarks of learned helplessness is trans-situationality. The dorsal raphe nucleus sends 5-HT projections to both the dorsal periaqueductal gray and to the amygdala, with 5-HT released in the dorsal periaqueductal gray inhibiting its function and 5-HT in the amygdala potentiating its function (see Graeff, Guimares, De Andrade, & Deakin, 1996 for review). Recall that escapable shock but not inescapable shock activates these labeled cells, as assessed by examining activation markers such as c-fos in these labeled neurons. So it seemed as if the subjects that received inescapable shock later behaved as if they had inhibited dorsal periaqueductal gray function and exaggerated amygdala/BNST function. Learned helplessness (after Martin Seligman) is a concept that holds that helplessness is learned and can be unlearned. Contributions of ERK signaling in the striatum to instrumental learning and performance. Seligman (Eds. Neurochemical basis of behavioral depression. IN produced a sustained rise in levels of extracellular 5-HT, while levels during ESC dropped rapidly as the subjects learned the controlling response. Otherwise, the measured change could be a simple consequence of the stressor per se. Weiss JM, Simson PE. The learning theories of that era held that organisms could only learn associations or pairings, for example a response paired with shock strengthened this association (acquisition) or a response paired with no shock weakened this association (extinction).

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