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1.  Rapid-Response Impulsivity: Definitions, Measurement Issues, and Clinical Implications 
Personality disorders  2015;6(2):168-181.
Impulsivity is a multi-faceted construct that is a core feature of multiple psychiatric conditions and personality disorders. However, progress in understanding and treating impulsivity in the context of these conditions is limited by a lack of precision and consistency in its definition and assessment. Rapid-response-impulsivity (RRI) represents a tendency toward immediate action that occurs with diminished forethought and is out of context with the present demands of the environment. Experts from the International Society for Research on Impulsivity (InSRI) met to discuss and evaluate RRI-measures in terms of reliability, sensitivity, and validity with the goal of helping researchers and clinicians make informed decisions about the use and interpretation of findings from RRI-measures. Their recommendations are described in this manuscript. Commonly-used clinical and preclinical RRI-tasks are described, and considerations are provided to guide task selection. Tasks measuring two conceptually and neurobiologically distinct types of RRI, “refraining from action initiation” (RAI) and “stopping an ongoing action” (SOA) are described. RAI and SOA-tasks capture distinct aspects of RRI that may relate to distinct clinical outcomes. The InSRI group recommends that: 1) selection of RRI-measures should be informed by careful consideration of the strengths, limitations, and practical considerations of the available measures; 2) researchers use both RAI and SOA tasks in RRI studies to allow for direct comparison of RRI types and examination of their associations with clinically relevant measures; and, 3) similar considerations should be made for human and non-human studies in an effort to harmonize and integrate pre-clinical and clinical research.
PMCID: PMC4476624  PMID: 25867840
impulsivity; response; attention; behavioral control; personality; assessment
2.  Recent Research on Impulsivity in Individuals With Drug Use and Mental Health Disorders: Implications for Alcoholism 
Alcohol misuse and dependence, and many of its accompanying psychological problems, are associated with heightened levels of impulsivity that both accelerate the development of clinically significant illness and complicate clinical outcome. This article reviews recent developments in our understanding of impulsivity as they relate to brain circuitry that might underlie these comorbid factors, focusing upon the clinical features of substance use (and dependence), bipolar disorder, and pathological gambling. Individuals who are affected by these disorders exhibit problems in several domains of impulsive behavior including deficient response or “motor” control, and the tolerance of prolonged delays prior to larger rewards at the expense of smaller rewards (“delay-discounting”). These populations, like alcoholic dependents, also exhibit impairments in risky decision-making that may reflect dysfunction of monoamine and catecholamine pathways. However, several areas of uncertainty exist including the specificity of impairments across disorders and the relationship between impulse control problems and altered evaluation of reward outcomes underlying observed impairments in action selection.
PMCID: PMC4777895  PMID: 20528825
Impulsivity; Substance Misuse; Alcohol; Pathological Gambling
3.  Missed losses loom larger than missed gains: Electrodermal reactivity to decision choices and outcomes in a gambling task 
Loss aversion is a defining characteristic of prospect theory, whereby responses are stronger to losses than to equivalently sized gains (Kahneman & Tversky Econometrica, 47, 263–291, 1979). By monitoring electrodermal activity (EDA) during a gambling task, in this study we examined physiological activity during risky decisions, as well as to both obtained (e.g., gains and losses) and counterfactual (e.g., narrowly missed gains and losses) outcomes. During the bet selection phase, EDA increased linearly with bet size, highlighting the role of somatic signals in decision-making under uncertainty in a task without any learning requirement. Outcome-related EDA scaled with the magnitudes of monetary wins and losses, and losses had a stronger impact on EDA than did equivalently sized wins. Narrowly missed wins (i.e., near-wins) and narrowly missed losses (i.e., near-losses) also evoked EDA responses, and the change of EDA as a function of the size of the missed outcome was modestly greater for near-losses than for near-wins, suggesting that near-losses have more impact on subjective value than do near-wins. Across individuals, the slope for choice-related EDA (as a function of bet size) correlated with the slope for outcome-related EDA as a function of both the obtained and counterfactual outcome magnitudes, and these correlations were stronger for loss and near-loss conditions than for win and near-win conditions. Taken together, these asymmetrical EDA patterns to objective wins and losses, as well as to near-wins and near-losses, provide a psychophysiological instantiation of the value function curve in prospect theory, which is steeper in the negative than in the positive domain.
PMCID: PMC4785217  PMID: 26667365
Prospect theory; Loss aversion; Near-miss; Arousal; Somatic marker hypothesis
4.  Role of Dopamine D2 Receptors in Human Reinforcement Learning 
Neuropsychopharmacology  2014;39(10):2366-2375.
Influential neurocomputational models emphasize dopamine (DA) as an electrophysiological and neurochemical correlate of reinforcement learning. However, evidence of a specific causal role of DA receptors in learning has been less forthcoming, especially in humans. Here we combine, in a between-subjects design, administration of a high dose of the selective DA D2/3-receptor antagonist sulpiride with genetic analysis of the DA D2 receptor in a behavioral study of reinforcement learning in a sample of 78 healthy male volunteers. In contrast to predictions of prevailing models emphasizing DA's pivotal role in learning via prediction errors, we found that sulpiride did not disrupt learning, but rather induced profound impairments in choice performance. The disruption was selective for stimuli indicating reward, whereas loss avoidance performance was unaffected. Effects were driven by volunteers with higher serum levels of the drug, and in those with genetically determined lower density of striatal DA D2 receptors. This is the clearest demonstration to date for a causal modulatory role of the DA D2 receptor in choice performance that might be distinct from learning. Our findings challenge current reward prediction error models of reinforcement learning, and suggest that classical animal models emphasizing a role of postsynaptic DA D2 receptors in motivational aspects of reinforcement learning may apply to humans as well.
PMCID: PMC4138746  PMID: 24713613
5.  Commentary on: Are we overpathologizing everyday life? A tenable blueprint for behavioral addiction research 
Journal of Behavioral Addictions  null;4(3):132-134.
Billieux et al. (2015) propose that the recent proliferation of behavioral addictions has been driven by deficiencies in the underlying research strategy. This commentary considers how pathological gambling (now termed gambling disorder) traversed these challenges to become the first recognized behavioral addiction in the DSM-5. Ironically, many similar issues continue to exist in research on gambling disorder, including question-marks over the validity of tolerance, heterogeneity in gambling motives, and the under-specification of neuroimaging biomarkers. Nevertheless, I contend that the case for gambling disorder as a behavioral addiction has been bolstered by the existence of clear and consistent functional impairment (primarily in the form of debt), coupled with the development of a public health approach that has given emphasis to product features (i.e. the structural characteristics of gambling forms) as much as individual dispositions (the ‘addictive personality’).
PMCID: PMC4627669  PMID: 26551898
pathological gambling; video games; addiction; tolerance; neuroimaging; structural features
6.  ‘Put Your Money Where Your Mouth Is!’: Effects of Streaks on Confidence and Betting in a Binary Choice Task 
Human choice under uncertainty is influenced by erroneous beliefs about randomness. In simple binary choice tasks, such as red/black predictions in roulette, long outcome runs (e.g. red, red, red) typically increase the tendency to predict the other outcome (i.e. black), an effect labeled the “gambler's fallacy.” In these settings, participants may also attend to streaks in their predictive performance. Winning and losing streaks are thought to affect decision confidence, although prior work indicates conflicting directions. Over three laboratory experiments involving red/black predictions in a sequential roulette task, we sought to identify the effects of outcome runs and winning/losing streaks upon color predictions, decision confidence and betting behavior. Experiments 1 (n = 40) and 3 (n = 40) obtained trial-by-trial confidence ratings, with a win/no win payoff and a no loss/loss payoff, respectively. Experiment 2 (n = 39) obtained a trial-by-trial bet amount on an equivalent scale. In each experiment, the gambler's fallacy was observed on choice behavior after color runs and, in experiment 2, on betting behavior after color runs. Feedback streaks exerted no reliable influence on confidence ratings, in either payoff condition. Betting behavior, on the other hand, increased as a function of losing streaks. The increase in betting on losing streaks is interpreted as a manifestation of loss chasing; these data help clarify the psychological mechanisms underlying loss chasing and caution against the use of betting measures (“post-decision wagering”) as a straightforward index of decision confidence. © 2014 The Authors. Journal of Behavioral Decision Making published by John Wiley & Sons Ltd.
PMCID: PMC4515090  PMID: 26236092
sequential biases; decision confidence; post-decision wagering; loss chasing
7.  Disappointment and regret enhance corrugator reactivity in a gambling task 
Psychophysiology  2014;52(4):518-523.
This study investigated how the corrugator and zygomaticus respond to decision outcomes (i.e., gains and losses). We used a gambling task in which participants were presented with obtained followed by non-obtained outcomes. Activity at the corrugator site was sensitive to decision outcomes, such that higher obtained losses (disappointment) and higher non-obtained gains (regret) both heightened corrugator reactivity. Activity at the zygomaticus site was not responsive to obtained or non-obtained outcomes, but did show sensitivity to emotional images in the same participants, in the form of a positive linear relationship with self-reported emotional valence. Corrugator activity was negatively related to emotional valence. The findings indicate the sensitivity of corrugator to objective decision outcomes and also counterfactual comparisons, highlighting the utility of facial electromyography in research on decision making and gambling behavior.
PMCID: PMC4510786  PMID: 25345723
Decision making; Gambling; Counterfactual thinking; Regret; Disappointment
8.  Near-wins and near-losses in gambling: A behavioral and facial EMG study 
Psychophysiology  2014;52(3):359-366.
This study investigated responses to near-wins (i.e., nonwin outcomes that were close to a major win, and their counterpart, near-losses (nonwin outcomes that are proximal to a major loss) in a decision-making task, measuring (a) luck ratings, (b) adjustment of bet amount, and (c) facial muscle reactivity at zygomaticus and corrugator sites. Compared to full-misses, near-wins decreased self-perceived luck and near-losses increased self-perceived luck, consistent with the effects of upward versus downward counterfactual thinking, respectively. Wins and losses both increased zygomaticus reactivity, and losses selectively enhanced corrugator reactivity. Near-wins heightened zygomaticus activity, but did not affect corrugator activity, thus showing a similar response pattern to actual wins. There were no significant facial EMG effects of near-losses. We infer that near-wins engender some appetitive processing, despite their objective nonwin status.
PMCID: PMC4510820  PMID: 25234840
Electromyography; Risk taking; Cognitive distortion; Near-miss; Gambling
9.  The cost of social punishment and high-lethality suicide attempts in the second half of life 
Psychology and aging  2014;29(1):84-94.
Age-related cognitive changes may contribute to impairments in making complex social decisions. Interpersonal conflict is a key factor behind suicidal behavior in old age, with suicidal motivations ranging from escape to revenge. Such conflicts may prove catastrophic for people prone to suicide, in part because of their tendency to make disadvantageous decisions. Yet, little is known about social decision-making in older suicidal individuals. We assessed economic bargaining behavior using the Ultimatum Game, where players decide whether to accept or punish (reject) unfair monetary offers from another player. Our sample included depressed older adults with a history of high-medical lethality suicide attempts, low- medical lethality suicide attempts, non-suicidal depressed older adults, and those with no psychiatric history, who served as control groups. Participants in all groups punished their counterparts in response to unfair offers. However, low-lethality attempters, non-suicidal depressed, and non-psychiatric controls punished less as the cost of punishment increased, accepting more unfair offers as the stakes grew large. High-lethality attempters did not adjust their choices based on stake magnitude, punishing unfair offers without regard to the cost. Two-thirds of the difference between the high-lethality attempters and non-psychiatric controls was explained by individual differences in fairness judgments: the comparison group judged offer fairness as a joint function of inequality and magnitude, whereas the high-lethality attempter participants judged offer fairness on the basis of inequality. In real life, high-lethality attempters' relative insensitivity to the cost of retaliation may lead to uncompromising, catastrophic responses to conflict.
PMCID: PMC4051315  PMID: 24660798
10.  Impaired executive function in contemplated and attempted suicide in late life 
Executive deficits may play an important role in late-life suicide. Yet, current evidence in this area is inconclusive and does not indicate whether these deficits are broadly associated with suicidal ideation or specific to suicidal behavior. This study examined global cognition and specifically executive function impairments as correlates of suicidal ideation and suicidal behavior in depressed older adults, with the goal of extending an earlier preliminary study.
Case-control study.
University-affiliated psychiatric hospital.
All were aged 60+: 83 depressed suicide attempters, 43 depressed individuals having suicidal ideation with a specific plan, 54 non-suicidal depressed participants, and 48 older adults with no history of psychiatric disorders.
Global cognitive function - Dementia Rating Scale (DRS), Executive function - Executive Interview (EXIT).
Both suicide attempters and suicide ideators performed worse than the two comparison groups on the EXIT, with no difference between suicide attempters and suicide ideators. On the DRS total score, as well as on Memory and Attention subscales, suicide attempters and ideators and non-suicidal depressed subjects performed similarly and were impaired relative to with non-psychiatric control subjects. Controlling for education, substance use disorders, and medication exposure did not affect group differences in performance on either the EXIT or DRS.
Executive deficits, captured with a brief instrument, are associated broadly with suicidal ideation in older depressed adults but do not appear to directly facilitate suicidal behavior. Our data are consistent with the idea that different vulnerabilities may operate at different stages in the suicidal process.
PMCID: PMC3516623  PMID: 23567385
suicide; cognitive; executive function; depression; aged
11.  Neurocognitive impairment in adolescent major depressive disorder: State vs. trait illness markers 
Journal of affective disorders  2011;133(3):625-632.
Current treatment outcomes of Major Depressive Disorder (MDD) in adolescents remain suboptimal. Discriminating between state and trait markers of MDD in adolescents would help identify markers that may guide choice of appropriate interventions and help improve longer-term outcome for individuals with the illness.
We compared neurocognitive performance in executive function, sustained attention and short-term memory in 20 adolescents with MDD in acute episode (MDDa), 20 previously depressed adolescents in remission (MDDr) and 17 healthy control participants (HC).
There was a group difference that emerged for executive function with increasing task difficulty (p = 0.033). MDDa showed impaired executive function, as measured by using more moves to solve 4-move problems on a forward planning task, relative to MDDr and HC (p = 0.01, d = 0.94 and p = 0.015, d = 0.77 respectively). MDDa showed more impulsivity as measured by lower response bias (B″) on a sustained attention task than both MDDr and HC (p = 0.01, d = 0.85 and p = 0.008, d = 0.49 respectively). Higher impulsivity was associated with more severe depression (r = −0.365, p = 0.022) and earlier age of onset of depression (r = 0.402, p = 0.012) and there was a trend for a correlation between more executive dysfunction and more severe depression (r = 0.301 p = 0.059) in MDDa and MDDr combined. The three groups did not differ significantly on short-term memory or target detection on the sustained attention task.
These results need to be replicated in the future with a larger sample size.
Executive dysfunction and impulsivity appear to be state-specific markers of MDD in adolescents that are related to depression severity and not present in remission.
PMCID: PMC4119611  PMID: 21620477
Depression; Adolescent; Neurocognition; Executive function; Impulsivity
12.  Striatal connectivity changes following gambling wins and near-misses: Associations with gambling severity 
NeuroImage : Clinical  2014;5:232-239.
Frontostriatal circuitry is implicated in the cognitive distortions associated with gambling behaviour. ‘Near-miss’ events, where unsuccessful outcomes are proximal to a jackpot win, recruit overlapping neural circuitry with actual monetary wins. Personal control over a gamble (e.g., via choice) is also known to increase confidence in one's chances of winning (the ‘illusion of control’).
Using psychophysiological interaction (PPI) analyses, we examined changes in functional connectivity as regular gamblers and non-gambling participants played a slot-machine game that delivered wins, near-misses and full-misses, and manipulated personal control. We focussed on connectivity with striatal seed regions, and associations with gambling severity, using voxel-wise regression.
For the interaction term of near-misses (versus full-misses) by personal choice (participant-chosen versus computer-chosen), ventral striatal connectivity with the insula, bilaterally, was positively correlated with gambling severity. In addition, some effects for the contrast of wins compared to all non-wins were observed at an uncorrected (p < .001) threshold: there was an overall increase in connectivity between the striatal seeds and left orbitofrontal cortex and posterior insula, and a negative correlation for gambling severity with the connectivity between the right ventral striatal seed and left anterior cingulate cortex.
These findings corroborate the ‘non-categorical’ nature of reward processing in gambling: near-misses and full-misses are objectively identical outcomes that are processed differentially. Ventral striatal connectivity with the insula correlated positively with gambling severity in the illusion of control contrast, which could be a risk factor for the cognitive distortions and loss-chasing that are characteristic of problem gambling.
PMCID: PMC4110887  PMID: 25068112
Gambling; Connectivity; fMRI; Reward; Near-miss; Addiction
13.  Pathological Choice: The Neuroscience of Gambling and Gambling Addiction 
The Journal of Neuroscience  2013;33(45):17617-17623.
Gambling is pertinent to neuroscience research for at least two reasons. First, gambling is a naturalistic and pervasive example of risky decision making, and thus gambling games can provide a paradigm for the investigation of human choice behavior and “irrationality.” Second, excessive gambling involvement (i.e., pathological gambling) is currently conceptualized as a behavioral addiction, and research on this condition may provide insights into addictive mechanisms in the absence of exogenous drug effects. 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.
PMCID: PMC3858640  PMID: 24198353
14.  Reward Signals, Attempted Suicide, and Impulsivity in Late-Life Depression 
JAMA psychiatry (Chicago, Ill.)  2013;70(10):10.1001/jamapsychiatry.2013.75.
Suicide can be viewed as an escape from unendurable punishment at the cost of any future rewards. Could faulty estimation of these outcomes predispose to suicidal behavior? In behavioral studies, many of those who have attempted suicide misestimate expected rewards on gambling and probabilistic learning tasks.
To describe the neural circuit abnormalities that underlie disadvantageous choices in people at risk for suicide and to relate these abnormalities to impulsivity, which is one of the components of vulnerability to suicide.
Case-control functional magnetic resonance imaging study of reward learning using a reinforcement learning model.
University hospital and outpatient clinic.
Fifty-three participants 60 years or older, including 15 depressed patients who had attempted suicide, 18 depressed patients who had never attempted suicide (depressed control subjects), and 20 psychiatrically healthy controls.
Components of the cortical blood oxygenation level–dependent response tracking expected and unpredicted rewards.
Depressed elderly participants displayed 2 distinct disruptions of control over reward-guided behavior. First, impulsivity and a history of suicide attempts (particularly poorly planned ones) were associated with a weakened expected reward signal in the paralimbic cortex, which in turn predicted the behavioral insensitivity to contingency change. Second, depression was associated with disrupted corticostriatothalamic encoding of unpredicted rewards, which in turn predicted the behavioral oversensitivity to punishment. These results were robust to the effects of possible brain damage from suicide attempts, depressive severity, co-occurring substance use and anxiety disorders, antidepressant and anticholinergic exposure, lifetime exposure to electroconvulsive therapy, vascular illness, and incipient dementia.
Altered paralimbic reward signals and impulsivity and/or carelessness may facilitate unplanned suicidal acts. This pattern, also seen in gambling and cocaine use, may reflect a primary deficit in the paralimbic cortex or in its mesolimbic input. The overreactivity to punishment in depression may be caused in part by a disruption of appetitive learning in the corticostriatothalamic circuits.
PMCID: PMC3859132  PMID: 23925710
16.  Risk-Sensitive Decision-Making in Patients with Posterior Parietal and Ventromedial Prefrontal Cortex Injury 
Successful choice under risk requires the integration of information about outcome probabilities and values and implicates a brain network including the ventromedial prefrontal cortex (vmPFC) and posterior parietal cortex (pPAR). Damage to the vmPFC is linked to poor decision-making and increased risk-taking. Electrophysiological and neuroimaging data implicate the pPAR in the processing of reward probability during choice, but the causal contribution of this area has not been established. We compared patients with lesions to the pPAR (n = 13), vmPFC (n = 13), and healthy volunteers (n = 22) on the Roulette Betting Task, a measure of risk-sensitive decision-making. Both lesion groups were impaired in adjusting their bets to the probability of winning. This impairment was correlated with the extent of pPAR, but not vmPFC, damage. In addition, the vmPFC group chose higher bets than healthy controls overall, an effect that correlated with lesion volume in the medial orbitofrontal cortex. Both lesion groups earned fewer points than healthy controls. The groups did not differ on 2 tasks assessing probabilistic reasoning outside of a risk-reward context. Our results demonstrate the causal involvement of both the pPAR and vmPFC in risk-sensitive choice and indicate distinguishable roles of these areas in probability processing and risk appetite.
PMCID: PMC4259274  PMID: 23926113
decision-making; lesion study; posterior parietal cortex; risk; ventromedial prefrontal cortex
17.  Restricting Temptations: Neural Mechanisms of Precommitment 
Neuron  2013;79(2):391-401.
Humans can resist temptations by exerting willpower, the effortful inhibition of impulses. But willpower can be disrupted by emotions and depleted over time. Luckily, humans can deploy alternative self-control strategies like precommitment, the voluntary restriction of access to temptations. Here, we examined the neural mechanisms of willpower and precommitment using fMRI. Behaviorally, precommitment facilitated choices for large delayed rewards, relative to willpower, especially in more impulsive individuals. While willpower was associated with activation in dorsolateral prefrontal cortex (DLPFC), posterior parietal cortex (PPC), and inferior frontal gyrus, precommitment engaged lateral frontopolar cortex (LFPC). During precommitment, LFPC showed increased functional connectivity with DLPFC and PPC, especially in more impulsive individuals, and the relationship between impulsivity and LFPC connectivity was mediated by value-related activation in ventromedial PFC. Our findings support a hierarchical model of self-control in which LFPC orchestrates precommitment by controlling action plans in more caudal prefrontal regions as a function of expected value.
•Precommitment is the voluntary restriction of access to temptations•Precommitment is a more effective self-control strategy than willpower•Precommitment engages LFPC; willpower engages DLPFC and PPC•During precommitment, LFPC increases connectivity with DLPFC and PPC
Crocket et al. show that during precommitment, which involves voluntarily restricting access to temptations, lateral frontopolar cortex is activated and increases functional connectivity with dorsolateral prefrontal and posterior parietal cortex—the same regions associated with actively resisting temptations.
PMCID: PMC3725418  PMID: 23889938
18.  The temptation of suicide: striatal gray matter, discounting of delayed rewards, and suicide attempts in late-life depression 
Psychological Medicine  2011;42(6):1203-1215.
Converging evidence implicates basal ganglia alterations in impulsivity and suicidal behavior. For example, D2/D3 agonists and subthalamic nucleus stimulation in Parkinson’s disease trigger impulse control disorders and possibly suicidal behavior. Further, suicidal behavior has been associated with structural basal ganglia abnormalities. Finally, low-lethality, unplanned suicide attempts are associated with increased discounting of delayed rewards, a behavior dependent upon the striatum. Thus, we tested whether, in late-life depression, changes in the basal ganglia were associated with suicide attempts and with increased delay discounting.
Fifty-two persons aged ≥60 underwent extensive clinical and cognitive characterization: 33 with major depression (13 suicide attempters [SA], 20 non-suicidal depressed elderly), and 19 non-depressed controls. Participants had high-resolution T1-weighted MPRAGE MRI scans. Basal ganglia gray matter voxel counts were estimated using atlas-based segmentation, with a highly-deformable automated algorithm. Discounting of delayed rewards was assessed using the Monetary Choice Questionnaire, and delay aversion with the Cambridge Gamble Task.
SA had lower putamen but not caudate or pallidum gray matter voxel counts, compared to the control groups. This difference persisted after accounting for substance use disorders and possible brain injury from suicide attempts. SA with lower putamen gray matter voxel counts displayed higher delay discounting on the MCQ, but not delay aversion on the CGT. Secondary analyses revealed that SA had lower voxel counts in associative and possibly ventral, but not sensorimotor striatum.
Our findings, while limited by small sample size and case-control design, suggest that striatal lesions could contribute to suicidal behavior by increasing impulsivity.
PMCID: PMC3368587  PMID: 21999930
suicide; basal ganglia; corpus striatum; globus pallidus; putamen; decision making; reward
19.  Fronto-striatal dysregulation in drug addiction and pathological gambling: Consistent inconsistencies?☆ 
NeuroImage : Clinical  2013;2:385-393.
Alterations in appetitive processing are central to the major psychological theories of addiction, with differential predictions made by the reward deficiency, incentive salience, and impulsivity hypotheses. Functional MRI has become the chief means of testing these predictions, with experiments reliably highlighting disturbances at the level of the striatum, medial prefrontal cortex, and affiliated regions. However, demonstrations of hypo-reactivity and hyper-reactivity of this circuitry in drug addicted groups are reported in approximately equal measure. Similar findings are echoed in the emergent neuroimaging literature on pathological gambling, which has recently witnessed a coming of age. The first aim of this article is to consider some of the methodological aspects of these experiments that could influence the observed direction of group-level effects, including the baseline condition, trial structure and timing, and the nature of the appetitive cues (drug-related, monetary, or primary rewards). The second aim is to highlight the conceptual traction that is offered by pathological gambling, as a model of a ‘toxicity free’ addiction and an illness where tasks of monetary reinforcement afford a more direct mapping to the abused commodity. Our conclusion is that relatively subtle decisions in task design appear capable of driving group differences in fronto-striatal circuitry in entirely opposing directions, even with tasks and task variants that look ostensibly similar. Differentiation between the psychological theories of addiction will require a greater breadth of experimental designs, with more research needed on processing of primary appetitive cues, aversive processing, and in vulnerable/at-risk groups.
•We outline the current psychological theories of addiction and their predictions.•We review recent fMRI literature of substance addictions and appetitive processing.•Reasons for opposing results (hyper- vs hypo-active reward regions) are discussed.•Recent fMRI findings of appetitive processing in pathological gambling are reviewed.•Pathological gambling is suggested as a prototypical addiction for imaging research.
PMCID: PMC3777686  PMID: 24179792
Addiction; Pathological gambling; fMRI; Ventral striatum; Appetitive processing
20.  Deterministic learning and attempted suicide among older depressed individuals: Cognitive assessment using the Wisconsin Card Sorting Task 
Journal of Psychiatric Research  2011;46(2):226-232.
Late-life suicide is an under-investigated public health problem. Among the putative vulnerabilities for this complex multifactorial behaviour are deficits in cognitive control, an ability to integrate and prioritize multiple cognitive processes in order to flexibly adapt behaviour and meet situational demands. We investigated cognitive control during rule learning in a complex and changing environment in older individuals with suicide attempts of varying lethality.
Ninety-three participants over the age of 60 (30 healthy controls, 29 depressed never suicidal, 20 low-lethality suicide attempters, 14 high-lethality suicide attempters) underwent structured clinical and cognitive assessments. Participants then completed the Wisconsin Card Sorting Test (WCST), a well-studied task of cognitive control during rule learning.
High-lethality attempters demonstrated a pattern of deficits involving poor conceptual reasoning, perseverative errors and total errors. Compared to low-lethality attempters and healthy controls, high-lethality attempters demonstrated poor conceptual reasoning, as well as increased rates of perseverative errors and total errors. Compared to non-suicidal depressed participants, high-lethality attempters also made more conceptual errors.
High-lethality suicide attempts among older people are associated with impaired cognitive control during rule learning as detected by the WCST. Our data suggest that impairment in cognitive control during rule learning may represent a vulnerability distinct from the impulsive diathesis, typically manifesting in young, low-lethality attempters. This vulnerability may contribute to the high incidence of serious or, often, fatal suicidal acts in old age.
PMCID: PMC3326354  PMID: 22024486
Suicide; Cognitive control; Cognitive function; Learning; Geriatrics; Depression
21.  Opioidergic and dopaminergic manipulation of gambling tendencies: a preliminary study in male recreational gamblers 
Gambling is characterized by cognitive distortions in the processing of chance and skill that are exacerbated in pathological gambling. Opioid and dopamine dysregulation is implicated in pathological gambling, but it is unclear whether these neurotransmitters modulate gambling distortions. The objective of the current study was to assess the effects of the opioid receptor antagonist naltrexone and the dopamine D2 receptor antagonist haloperidol on gambling behavior. Male recreational gamblers (n = 62) were assigned to receive single oral doses of naltrexone 50 mg, haloperidol 2 mg or placebo, in a parallel-groups design. At 2.5 h post-dosing, participants completed a slot machine task to elicit monetary wins, “near-misses,” and a manipulation of personal choice, and a roulette game to elicit two biases in sequential processing, the gambler's fallacy and the hot hand belief. Psychophysiological responses (electrodermal activity and heart rate) were taken during the slot machine task, and plasma prolactin increase was assessed. The tasks successfully induced the gambling effects of interest. Some of these effects differed across treatment groups, although the direction of effect was not in line with our predictions. Differences were driven by the naltrexone group, which displayed a greater physiological response to wins, and marginally higher confidence ratings on winning streaks. Prolactin levels increased in the naltrexone group, but did not differ between haloperidol and placebo, implying that naltrexone but not haloperidol may have been functionally active at these doses. Our results support opioid modulation of cognition during gambling-like tasks, but did not support the more specific hypothesis that naltrexone may act to ameliorate cognitive distortions.
PMCID: PMC3791382  PMID: 24109443
naltrexone; haloperidol; pathological gambling; addiction; reward; motivation; decision-making; psychophysiology
22.  Striatal dopamine D2/D3 receptor binding in pathological gambling is correlated with mood-related impulsivity 
Neuroimage  2012;63(1):40-46.
Pathological gambling (PG) is a behavioural addiction associated with elevated impulsivity and suspected dopamine dysregulation. Reduced striatal dopamine D2/D3 receptor availability has been reported in drug addiction, and may constitute a premorbid vulnerability marker for addictive disorders. The aim of the present study was to assess striatal dopamine D2/D3 receptor availability in PG, and its association with trait impulsivity. Males with PG (n = 9) and male healthy controls (n = 9) underwent [11C]-raclopride positron emission tomography imaging and completed the UPPS-P impulsivity scale. There was no significant difference between groups in striatal dopamine D2/D3 receptor availability, in contrast to previous reports in drug addiction. However, mood-related impulsivity (‘Urgency’) was negatively correlated with [11C]-raclopride binding potentials in the PG group. The absence of a group difference in striatal dopamine binding implies a distinction between behavioural addictions and drug addictions. Nevertheless, our data indicate heterogeneity in dopamine receptor availability in disordered gambling, such that individuals with high mood-related impulsivity may show differential benefits from dopamine-based medications.
► Assessed 11C-raclopride binding in pathological gambling, a putative behavioral addiction. ► No group difference in striatal dopamine binding from healthy controls. ► Dopamine binding negatively correlated with mood-related impulsivity (‘Urgency’).
PMCID: PMC3438449  PMID: 22776462
Gambling; Impulsivity; Dopamine; Neuroimaging; Addiction; Striatum
23.  Serotonin Modulates the Effects of Pavlovian Aversive Predictions on Response Vigor 
Neuropsychopharmacology  2012;37(10):2244-2252.
Updated theoretical accounts of the role of serotonin (5-HT) in motivation propose that 5-HT operates at the intersection of aversion and inhibition, promoting withdrawal in the face of aversive predictions. However, the specific cognitive mechanisms through which 5-HT modulates withdrawal behavior remain poorly understood. Behavioral inhibition in response to punishments reflects at least two concurrent processes: instrumental aversive predictions linking stimuli, responses, and punishments, and Pavlovian aversive predictions linking stimuli and punishments irrespective of response. In the current study, we examined to what extent 5-HT modulates the impact of instrumental vs Pavlovian aversive predictions on behavioral inhibition. We used acute tryptophan depletion to lower central 5-HT levels in healthy volunteers, and observed behavior in a novel task designed to measure the influence of Pavlovian and instrumental aversive predictions on choice (response bias) and response vigor (response latencies). After placebo treatment, participants were biased against responding on the button that led to punishment, and they were slower to respond in a punished context, relative to a non-punished context. Specifically, participants slowed their responses in the presence of stimuli predictive of punishments. Tryptophan depletion removed the bias against responding on the punished button, and abolished slowing in the presence of punished stimuli, irrespective of response. We suggest that this set of results can be explained by a role for 5-HT in Pavlovian aversive predictions. These findings suggest additional specificity for the influence of 5-HT on aversively motivated behavioral inhibition and extend recent models of the role of 5-HT in aversive predictions.
PMCID: PMC3422489  PMID: 22643930
serotonin; aversion; inhibition; tryptophan; Pavlovian; instrumental; Serotonin; Cognition; Psychopharmacology; Behavioral Science; aversion; inhibition; tryptophan; Pavlovian; instrumental
24.  Lethal Forethought: Delayed Reward Discounting Differentiates High- and Low-Lethality Suicide Attempts in Old Age 
Biological psychiatry  2011;70(2):138-144.
The decision to commit suicide may be impulsive, but lethal suicidal acts often involve planning and forethought. People who attempt suicide make disadvantageous decisions in other contexts, but nothing is known about the way they decide about the future. Can the willingness to postpone future gratification differentiate between individuals prone to serious, premeditated and less serious, unplanned suicidal acts?
Four groups of depressed participants aged 60+ made choices between smaller immediate and larger delayed monetary rewards: 15 who made high-lethality suicide attempts, 14 who made low-lethality suicide attempts, 12 who seriously contemplated suicide, and 42 people with depression but no history of suicidal thoughts. The reference group was 31 psychiatrically healthy elders.
Individuals who had made low-lethality attempts displayed an exaggerated preference for immediate rewards compared to non-suicidal depressed and healthy controls. Those who had carried out high-lethality suicide attempts were more willing to delay future rewards, compared to low-lethality attempters. Better planned suicide attempts were also associated with willingness to wait for larger rewards. These effects were unchanged after accounting for education, global cognitive function, substance use disorders, psychotropic medications, and possible brain injury from attempts. Discount rates were correlated with having debt but were not significantly associated with income, hopelessness, depressive severity, premorbid IQ, age at first attempt, or choice of violent means.
While clinicians often focus on impulsivity in patients at risk for suicide, these data suggest that identifying biological characteristics and treatments for non-impulsive suicidal older people may be even more important.
PMCID: PMC3125431  PMID: 21329911
suicide; cognition; decision making; reward; depressive disorder; aged; time perception; executive function; choice behavior
25.  Gut feelings and the reaction to perceived inequity: The interplay between bodily responses, regulation, and perception shapes the rejection of unfair offers on the ultimatum game 
It has been robustly demonstrated using the ultimatum game (UG) that individuals frequently reject unfair financial offers even if this results in a personal cost. One influential hypothesis for these rejections is that they reflect an emotional reaction to unfairness that overrides purely economic decision processes. In the present study, we examined whether the interplay between bodily responses, bodily regulation, and bodily perception (“interoception”) contributes to emotionally driven rejection behavior on the UG. Offering support for bodily feedback theories, interoceptive accuracy moderated the relationship between changes in electrodermal activity to proposals and the behavioral rejection of such offers. Larger electrodermal responses to rejected relative to accepted offers predicted greater rejection in those with accurate interoception but were unrelated to rejection in those with poor interoception. Although cardiovascular responses during the offer period were unrelated to rejection rates, greater resting heart rate variability (linked to trait emotion regulation capacity) predicted reduced rejection rates of offers. These findings help clarify individual differences in reactions to perceived unfairness, support previous emotion regulation deficit accounts of rejection behavior, and suggest that the perception and regulation of bodily based emotional biasing signals (“gut feelings”) partly shape financial decision making on the UG.
PMCID: PMC3400033  PMID: 22618636
Decision-making; Embodied cognition; Emotion

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