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Pathological Choice: The Neuroscience of Gambling and Gambling Addiction

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Postby Mecage В» 17.03.2020

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Author contributions: L. Gambling is pertinent to neuroscience research for at least two reasons. This article is a summary of topics covered in a Society for Neuroscience minisymposium, focusing on recent advances in understanding the neural basis of gambling behavior, including translational findings in rodents and nonhuman primates, which have begun to delineate neural circuitry and neurochemistry involved.

Examining their underlying neural mechanisms is naturally relevant to the emergent discipline of neuroeconomics. Gambling also has a more insidious side. Pathological gambling was first recognized as a psychiatric disorder in and was grouped initially in the Impulse Control Disorders.

An international program of research over the past decade has revealed multiple similarities between pathological gambling and the substance use disorders, including neurobiological overlap Petry, , Leeman and Potenza, The current article aims to provide a concise overview of recent developments in our understanding of decision making during gambling and the relevance of these processes to problem gambling for comprehensive overviews, see van Holst et al.

We begin by describing some emerging methods for probing gambling decisions, highlighting translational models, behavioral economic tasks, and cognitive distortions associated with gambling Fig. We then consider the underlying neural mechanisms, distinguishing neurochemical substrates and neuroanatomy. Schematic overview showing the emerging methods for modeling gambling decisions and the associated neural circuitry.

The list is not intended as comprehensive but highlights the core themes covered in this review. Given that the calculation of risk versus reward trade-offs is inherent in numerous aspects of real-world choice and foraging behavior, it should be unsurprising that laboratory animals are capable of performing decision-making tasks that resemble gambling.

Recent work has aimed to model gambling decisions in rats using operant behavioral tasks derived from the established probes of choice behavior in human neuropsychology and cognitive psychology. One widely used human test is the Iowa Gambling Task Bechara et al. In humans, this task involves a series of choices between four decks of cards that offer gains and losses of varying amounts of money. In the rat Gambling Task Zeeb et al.

Like the human version, the two apertures that offer larger rewards are also associated with longer and more frequent time-outs, and most rats learn to avoid these tempting options to maximize their sugar pellet profits over the duration of the task.

The key decision here is probabilistic and the task should not be confused with temporal discounting. Postacquisition lesions to BLA skewed rats' preference toward the high-risk high-reward options, matching the observation that amygdala damage leads to disadvantageous choice in the Iowa Gambling Task Bechara et al. If BLA lesions were made before task acquisition, animals struggled to develop the optimal strategy and correctly discriminate between the options.

Lesions to the orbitofrontal cortex OFC impaired acquisition of the rodent task in an identical manner but did not affect performance if the lesions were implemented after animals had learned the correct strategy. Such data support the suggestion that the classic disruption of everyday decision making associated with ventromedial prefrontal cortex lesions may stem from a difficulty in learning the optimal strategy, rather than an increase in preference for risky outcomes per se Bechara et al.

Moreover, the similarity between the effects of BLA and OFC lesions on task acquisition suggested that these two areas work together to promote development of the optimal strategy, a hypothesis recently confirmed using a functional disconnection procedure Zeeb and Winstanley, Hence, similar brain regions appear to be involved in guiding decision making under uncertainty in both rats and humans.

Prefrontal connectivity with the striatum is also implicated in choice behavior. Contemporary hypotheses of frontostriatal function emphasize a primary role in either action selection or reinforcement learning, both of which are likely important in substance addiction and behavioral addictions. To differentiate these elements, Seo et al. While the animals performed this task, neural activity was monitored simultaneously in anatomically connected regions of lateral prefrontal cortex LPFC; caudal area 46 and the dorsal striatum DS, primarily the anterior caudate nucleus.

A larger fraction of LPFC neurons represented selected actions, independent of how they were selected. In the perceptual inference condition, the LPFC representation of the selected action preceded the DS representation of the selected action, whereas in the reinforcement learning condition, both structures represented the actions up to ms before they were executed, with no clear temporal ordering.

Additionally, DS more often represented the value of the selected action when it was selected using both perceptual inference and reinforcement learning. Thus, a hypothesis that the DS was important for action selection was not supported, but DS did often represent action values, when driven by either reinforcement learning or perceptual inference. LPFC, by contrast, appears to play a dominant role in representing and selecting actions, particularly when the selection is based on perceptual inference.

Behavioral economic aims to decompose the processes of option valuation into simple components that can be quantified with discrete parameters Schonberg et al. Prospect theory PT remains the most influential of these accounts because of its ability to describe a range of common behaviors and deviations from normative expected value theory Kahneman and Tversky, For example, subjects typically reject mixed gambles that offer a 50—50 chance of winning or losing a given amount of money.

Loss aversion may be underpinned by value computations in the ventral striatum and amygdala Tom et al. In addition to this asymmetry between gains and losses, PT describes a value function for gains that is concave, contrasting with a value function for losses that is convex. Although recent work has demonstrated impaired processing of loss information Brevers et al. Behaviorally, the overestimation of small probabilities may contribute to the attractiveness of gambles, such as a lottery Trepel et al.

Ligneul et al. As expected, the results revealed elevated risk taking in gamblers compared with nongambling controls; however, this behavior was not linked to a specific distortion of small probabilities but rather to a general overweighting across the entire probability range.

Similar approaches using the discounting framework have demonstrated fine alterations of value representations in the ventral striatum in pathological gamblers Miedl et al. In addition to the computational characterization of gambling offered by behavioral economics, psychological models of gambling have additionally highlighted the central role of cognitive distortions during gambling.

These distortions refer to how the gambler thinks about randomness, chance, and skill Ladouceur and Walker, ; Clark, and foster an inappropriately high expectation of winning during the game. A number of specific biases have been described, and these cognitions can be effectively targeted as one element of psychotherapy for pathological gambling Fortune and Goodie, Arguably, the most classic distortion is the gambler's fallacy, which is a bias in the processing of random sequences.

In this compelling example, the expectancy of a certain event e. The phenomenon occurs across many situations, including casino gambling Sundali and Croson, , but also stock investment Johnson et al. It is widely viewed as arising from the representativeness heuristic, the belief that a short segment of a random sequence should reflect the overall distribution Rabin, ; Ayton and Fischer, Illusory control can be fostered by a various psychological features of games, such as the involvement of a choice e.

A recent study using a contingency judgment task from the associative learning literature found that pathological gamblers displayed a greater tendency to overestimate their control of positive outcomes than nongambling participants Orgaz et al. Using a slot machine task that delivered occasional jackpot wins, near misses where the reels landed adjacent to a win were associated with higher self-reported motivations to gamble than full-miss outcomes, despite their objective equivalence as nonwins Clark et al.

To date, most of the research on these distortions has been in healthy samples; and although questionnaire measures, such as the Gambling Related Cognitions Scale Raylu and Oei, , indicate a clear increase in the overall propensity toward these erroneous beliefs in pathological gamblers Michalczuk et al.

Dopamine has been a prime candidate for investigation of neurochemical abnormalities in pathological gamblers, given its established roles in both drug addiction and rewarded behavior. In patients with Parkinson's disease, sudden onset gambling can be observed, alongside other reward-driven behaviors, including compulsive shopping and hypersexuality, as a side effect of dopamine agonist medications Ambermoon et al.

Ongoing work is using alternative PET tracers that offer advantages over [ 11 C]-raclopride. The D 3 receptor subtype is localized to limbic circuitry and implicated in drug self-administration and relapse behavior in preclinical models Heidbreder et al.

This association suggests that D 3 expression is relevant to symptom severity in problem gambling, and as an addiction phenotype, it may be a useful marker for risk. Preliminary work has also begun to examine dopamine release in pathological gamblers, with some provocative early findings.

Payer, I. Boileau, D. Lobo, B. Chugani, A. Behzadi, A. Wilson, S. Kish, S. Houle, M. Zack, unpublished observations. This result effect is echoed in two further experiments examining task-related changes in [ 11 C]-raclopride binding in pathological gamblers, where higher levels of dopamine release were correlated with greater subjective excitement Linnet et al. Notably, the available data in drug addiction show blunted dopamine release in response to psychostimulant administration Volkow et al.

The extent to which these discrepancies reflect etiological differences between substance and behavioral addictions, or the masking of incentive sensitization processes via drug-induced depletion of dopamine stores Robinson and Berridge, , is a key question in ongoing research.

Rodent models have also provided a means of examining the neurochemistry of gambling, implicating dopamine and serotonin influences. In light of the effects of dopamine agonist medications in Parkinson's disease, it is notable that administration of selective D 2 agonists did not affect choice behavior on the rat Gambling Task Zeeb et al. Selective dopamine antagonists did not block the effects of amphetamine on choice, even though such agents did attenuate amphetamine-induced increases in motor impulsivity Zeeb et al.

Furthermore, amphetamine's effects were not mimicked by dopamine, 5-HT, or noradrenaline reuptake inhibitors but were reproduced by different combinations of these drugs Baarendse et al.

Such results imply concurrent regulation of choice behavior on the rat Gambling Task by multiple monoaminergic systems, consistent with human data Rogers, In this task, the rats choose between two options of equivalent value, one of which delivers a guaranteed reward, and the other offers either double that reward or nothing, with odds.

The reward size varies over the session from 1 to 3 pellets. Critically, such a behavioral shift confers no advantage in terms of reward earned and may be considered irrational in a similar vein to the framing effects observed in human choice under risk. Functional neuroimaging studies have also contributed much to our understanding of appetitive processing in pathological gamblers and provide data that complement the investigations of dopamine transmission Schott et al.

This observation can be interpreted in terms of the reward deficiency hypothesis Comings and Blum, , consistent with the PET evidence reviewed above indicating reduced dopamine receptor levels in addiction.

However, other recent studies have described increased, rather than decreased, responses to monetary rewards in the same population Hewig et al. One means of resolving these discrepancies is to consider the sensitivity to nonmonetary i.

Using an incentive delay protocol involving both monetary and visual erotic rewards, pathological gamblers showed a markedly decreased response to the erotic cues, compared with monetary cues, in the ventral striatum Sescousse et al. This differential response was correlated with the severity of gambling symptoms and accompanied by a similarly reduced behavioral motivation for erotic rewards.

Comparable designs indicate blunted brain responses to non—drug-related cues in drug-addicted groups Goldstein et al. These findings suggest that the key variable of interest may be the differential response to monetary or drug rewards versus other primary appetitive cues, rather than the response to money or drugs per se.

Experiments on gambling-related cognitive distortions also implicate reward-related circuitry, as well as the interactions with regions responsible for top-down cognitive control.

Specifically, the gambler's fallacy appears to arise from an imbalance between cognitive and emotional decision making mechanisms in the brain Shiv et al. Using a card guessing task to capture subjects' tendency to predict the break of a streak as it continued a signal of the gambler's fallacy , enhanced neural responses in left LPFC were observed to outcomes that were followed by a gambler's fallacy switch Xue et al.

A follow-up experiment applied anodal transcranial direct current stimulation, a procedure known to enhance cortical excitability and cerebral perfusion Stagg et al. Thus, the gambler's fallacy seems to be associated with 1 weak function in the affective decision making system and 2 strong function in the LPFC cognitive control system Xue et al. In contrast to these cortical responses, the robust striatal activations seen in response to monetary wins are not evidently modulated by the psychological context that characterizes these gambling distortions.

For example, the striatal responses to winning outcomes did not differ between the first win in a streak, compared with the fourth successive win Akitsuki et al. In a study investigating the illusion of control, striatal activity did not differ between choice and no-choice conditions, even though perceived control did enhance subjective confidence Kool et al.

However, both distortions appear to be coded in higher cortical regions. Functional imaging results revealed that the decision-related activation in the lateral and medial PFC was significantly modulated by both agency and previous outcome and that these effects were further predicted by the trait-like disposition to attribute negative events externally.

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Postby Tojatilar В» 17.03.2020

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