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1.  Dissociable neural representations of future reward magnitude and delay during temporal discounting 
NeuroImage  2008;45(1):143-150.
In temporal discounting, individuals often prefer smaller immediate rewards to larger delayed rewards, implying a trade off between the magnitude and delay of future rewards. While recent functional magnetic resonance imaging (fMRI) investigations of temporal discounting have generated conflicting findings, no studies have focused on whether distinct neural substrates respond to the magnitude and delay of future rewards. Combining a novel, temporally distributed discounting task with event-related fMRI, we found that while nucleus accumbens (NAcc), mesial prefrontal cortical (MPFC), and posterior cingulate cortical (PCC) activation positively correlated with future reward magnitude, dorsolateral prefrontal cortical (DLPFC) and posterior parietal cortical (PPC) activation negatively correlated with future reward delay. Further, more impulsive individuals showed diminished NAcc activation to the magnitude of future rewards and greater deactivations to delays of future rewards in the MPFC, DLPFC, and PPC. These findings suggest that while mesolimbic dopamine projection regions show greater sensitivity to the magnitude of future rewards, lateral cortical regions show greater (negative) sensitivity to the delay of future rewards, potentially reconciling different neural accounts of temporal discounting.
PMCID: PMC2685201  PMID: 19071223
temporal discounting; reward; accumbens; prefrontal; individual differences; FMRI; decision; human
2.  Neural and Behavioral Mechanisms of Impulsive Choice in Alcohol Use Disorder 
Alcohol dependence has repeatedly been associated with impulsive choice, or the inability to choose large delayed rewards over smaller, but more immediate rewards. However, the neural basis of impulsive choice in alcohol use disorders is not well understood.
One hundred fifty one individuals with a range of alcohol use from social drinking to severe alcohol dependence completed a delay discounting task while undergoing functional magnetic resonance imaging (fMRI). Participants received customized trials designed to ensure an approximately equivalent number of immediate responses.
Delaying gratification recruited regions involved in cognitive control, conflict monitoring, and the interpretation of somatic states. Individuals with more severe alcohol use problems showed increased discounting of delayed rewards and greater activation in several regions including supplementary motor area, insula/orbitofrontal cortex, inferior frontal gyrus, and the precuneus.
These results suggest that impulsive choice in alcohol dependence is the result of functional anomolies in widely distributed, but interconnected brain regions involved in cognitive and emotional control. Further, our results suggest that the neural mechanisms of impulsive choice in alcohol use disorders both overlaps with that observed in previous studies, and shows that individuals with alcohol use disorders recruit additional mechanisms when making intertemporal choices.
PMCID: PMC3117198  PMID: 21676001
Delay discounting; intertemporal choice; impulsivity; delayed gratification
3.  The neuroeconomics of nicotine dependence: A preliminary study of delay discounting of monetary and cigarette rewards in smokers using fMRI 
Psychiatry research  2012;202(1):20-29.
Neuroeconomics integrates behavioral economics and cognitive neuroscience to understand the neurobiological basis for normative and maladaptive decision making. Delay discounting is a behavioral economic index of impulsivity that reflects capacity to delay gratification and has been consistently associated with nicotine dependence. This preliminary study used functional magnetic resonance imaging to examine delay discounting for money and cigarette rewards in 13 nicotine dependent adults. Significant differences between preferences for smaller immediate rewards and larger delayed rewards were evident in a number of regions of interest (ROIs), including the medial prefrontal cortex, anterior insular cortex, middle temporal gyrus, middle frontal gyrus, and cingulate gyrus. Significant differences between money and cigarette rewards were generally lateralized, with cigarette choices associated with left hemisphere activation and money choices associated with right hemisphere activation. Specific ROI differences included the posterior parietal cortex, medial and middle frontal gyrus, ventral striatum, temporoparietal cortex, and angular gyrus. Impulsivity as measured by behavioral choices was significantly associated with both individual ROIs and a combined ROI model. These findings provide initial evidence in support of applying a neuroeconomic approach to understanding nicotine dependence.
PMCID: PMC3530421  PMID: 22633679
Nicotine dependence; smoking; tobacco; behavioral economics; neuroeconomics; delay discounting; impulsivity
4.  Delay Discounting and Frontostriatal Fiber Tracts: A Combined DTI and MTR Study on Impulsive Choices in Healthy Young Adults 
Cerebral Cortex (New York, NY)  2012;23(7):1695-1702.
Delay discounting, a measure of impulsive choice, has been associated with decreased control of the prefrontal cortex over striatum responses. The anatomical connectivity between both brain regions in delaying gratification remains unknown. Here, we investigate whether the quality of frontostriatal (FS) white matter tracts can predict individual differences in delay-discounting behavior. We use tract-based diffusion tensor imaging and magnetization transfer imaging to measure the microstructural properties of FS fiber tracts in 40 healthy young adults (from 18 to 25 years). We additionally explored whether internal sex hormone levels affect the integrity of FS tracts, based on the hypothesis that sex hormones modulate axonal density within prefrontal dopaminergic circuits. We calculated fractional anisotropy (FA), mean diffusivity (MD), longitudinal diffusivity, radial diffusivity (RD), and magnetization transfer ratio (MTR), a putative measure of myelination, for the FS tract. Results showed that lower integrity within the FS tract (higher MD and RD and lower FA), predicts faster discounting in both sexes. MTR was unrelated to delay-discounting performance. In addition, testosterone levels in males were associated with a lower integrity (higher RD) within the FS tract. Our study provides support for the hypothesis that enhanced structural integrity of white matter fiber bundles between prefrontal and striatal brain areas is associated with better impulse control.
PMCID: PMC3673180  PMID: 22693341
delay discounting; DTI; frontostriatal tracts; impulsivity; testosterone
5.  Frontal Hyperconnectivity Related to Discounting and Reversal Learning in Cocaine Subjects 
Biological psychiatry  2011;69(11):1117-1123.
Functional neuroimaging studies suggest that chronic cocaine use is associated with frontal lobe abnormalities. Functional connectivity (FC) alterations of cocaine dependent individuals (CD), however, are not yet clear. This is the first study to our knowledge that examines resting FC of anterior cingulate cortex (ACC) in CD. Because ACC is known to integrate inputs from different brain regions to regulate behavior, we hypothesize that CD will have connectivity abnormalities in ACC networks. In addition, we hypothesized that abnormalities would be associated with poor performance in delayed discounting and reversal learning tasks.
Resting functional magnetic resonance imaging data were collected to look for FC differences between twenty-seven cocaine dependent individuals (CD) (5 females, age: M=39.73, SD=6.14) and twenty-four controls (5 females, age: M=39.76, SD = 7.09). Participants were assessed with delayed discounting and reversal learning tasks. Using seed-based FC measures, we examined FC in CD and controls within five ACC connectivity networks with seeds in subgenual, caudal, dorsal, rostral, and perigenual ACC.
CD showed increased FC within the perigenual ACC network in left middle frontal gyrus, ACC and middle temporal gyrus when compared to controls. FC abnormalities were significantly positively correlated with task performance in delayed discounting and reversal learning tasks in CD.
The present study shows that participants with chronic cocaine-dependency have hyperconnectivity within an ACC network known to be involved in social processing and mentalizing. In addition, FC abnormalities found in CD were associated with difficulties with delay rewards and slower adaptive learning.
PMCID: PMC3090521  PMID: 21371689
cocaine; functional connectivity; anterior cingulate; delayed discount; reversal learning; frontal
6.  How Many Impulsivities? A Discounting Perspective 
People discount the value of delayed and uncertain outcomes, and how steeply individuals discount is thought to reflect how impulsive they are. From this perspective, steep discounting of delayed outcomes (which fails to maximize long-term welfare) and shallow discounting of probabilistic outcomes (which fails to adequately take risk into account) reflect the same trait of impulsivity. Despite the fact that a hyperboloid function describes the discounting of both delayed and probabilistic outcomes, there is considerable evidence that the two kinds of discounting involve different processes as well as separate impulsivity traits. Several manipulations differentially affect delay and probability discounting, and correlational studies show that how steeply one discounts delayed rewards is relatively independent of how steeply one discounts probabilistic rewards. Moreover, people’s discounting of delayed money and health outcomes are uncorrelated as are discounting of real, consumable rewards and hypothetical money. These results suggest that even within delay discounting, there may be multiple ‘impulsivities,’ each of which may be important for understanding a different aspect of decision making. Taken together, the pattern of findings reviewed here argues for a more nuanced view of impulsivity than that which is usually assumed in discounting research.
PMCID: PMC3893105  PMID: 23344985
discounting; impulsivity; traits; delay; probability
7.  Greater Impulsivity is Associated with Decreased Brain Activation in Obese Women during a Delay Discounting Task 
Brain imaging and behavior  2013;7(2):116-128.
Impulsivity and poor inhibitory control are associated with higher rates of delay discounting (DD) or a greater preference for smaller, more immediate rewards at the expense of larger, but delayed rewards. Of the many functional magnetic resonance imaging (fMRI) studies of DD, few have investigated the correlation between individual differences in DD rate and brain activation related to DD trial difficulty, with difficult DD trials expected to activate putative executive function brain areas involved in impulse control. In the current study, we correlated patterns of brain activation as measured by fMRI during difficult vs. easy trials of a DD task with DD rate (k) in obese women. Difficulty was defined by how much a reward choice deviated from an individual’s ‘indifference point’, or the point where the subjective preference for an immediate and a delayed reward was approximately equivalent. We found that greater delay discounting was correlated with less modulation of activation in putative executive function brain areas, such as the middle and superior frontal gyri and inferior parietal lobule, in response to difficult compared to easy DD trials. These results support the suggestion that increased impulsivity is associated with deficient functioning of executive function areas of the brain.
PMCID: PMC3561478  PMID: 22948956
impulsivity; inhibitory control; executive function; delay discounting; intertemporal
8.  Cortical activation during delay discounting in abstinent methamphetamine dependent individuals 
Psychopharmacology  2008;201(2):183-193.
Methamphetamine (MA)-dependent individuals prefer smaller immediate over larger delayed rewards in delay discounting (DD) tasks. Human and animal data implicate ventral (amygdala, ventral striatum, ventrolateral prefrontal cortex insula) and dorsal (dorsolateral prefrontal cortex, dorsal anterior cingulate cortex and posterior parietal cortex) systems in DD decisions. The ventral system is hypothesized to respond to the salience and immediacy of rewards while the dorsal system is implicated in the process of comparison and choice.
We used functional Magnetic Resonance Imaging to probe the neural correlates of DD in 19 recently abstinent MA-dependent patients and 17 age- and gender-matched controls.
Hard DD choices were associated with greatest activation in bilateral middle cingulate, posterior parietal cortex (PPC), and the right rostral insula. Control subjects showed more activation than MA patients bilaterally in the precuneus and in the right caudate nucleus, anterior cingulate cortex (ACC), and dorsolateral prefrontal cortex (DLPFC). Magnitude of discounting was correlated with activity in the amygdala, DLPFC, posterior cingulate cortex and PPC.
Our findings were consistent with a model wherein dorsal cognitive systems modulate the neural response of ventral regions. Patients addicted to MA, who strongly prefer smaller immediate over larger delayed rewards, activate the dorsal cognitive control system in order to overcome their preference. Activation of the amygdala during choice of delayed rewards was associated with a greater degree of discounting, suggesting that heavily discounting MA-dependent individuals may be more responsive to the negative salience of delayed rewards than controls.
PMCID: PMC2835463  PMID: 18685833
Methamphetamine; Delay discounting; Brain imaging
9.  Dissociable Brain Signatures of Choice Conflict and Immediate Reward Preferences in Alcohol Use Disorders 
Addiction biology  2012;10.1111/adb.12017.
Impulsive delayed reward discounting (DRD) is an important behavioral process in alcohol use disorders (AUDs), reflecting incapacity to delay gratification. Recent work in neuroeconomics has begun to unravel the neural mechanisms supporting DRD, but applications of neuroeconomics in relation to AUDs have been limited. This study examined the neural mechanisms of DRD preferences in AUDs, with emphasis on dissociating activation patterns based on DRD choice type and level of cognitive conflict. Heavy drinking adult males with (n = 13) and without (n = 12) a diagnosis of an AUD completed a monetary DRD task during a functional magnetic resonance imaging scan. Participant responses were coded based on choice type (impulsive vs. restrained) and level of cognitive conflict (easy vs. hard). AUD+ participants exhibited significantly more impulsive DRD decision-making. Significant activation during DRD was found in several decision-making regions, including dorsolateral prefrontal cortex (DLPFC), insula, posterior parietal cortex (PPC), and posterior cingulate. An axis of cognitive conflict was also observed, with hard choices associated with anterior cingulate cortex and easy choices associated with activation in supplementary motor area. AUD+ individuals exhibited significant hyperactivity in regions associated with cognitive control (DLPFC) and prospective thought (PPC) and exhibited less task-related deactivation of areas associated with the brain's default network during DRD decisions. This study provides further clarification of the brain systems supporting DRD in general and in relation to AUDs.
PMCID: PMC3871988  PMID: 23231650
Alcohol use disorders; delay discounting; neuroeconomics
The present study explored the delay discounting of future and past monetary rewards by pathological gamblers. Using a multiple baseline design, following repeated exposure to choices between smaller immediate and larger delayed consequences, participants completed a relational responding task that attempted to alter the psychological functions of irrelevant stimuli and to affect subsequent delay discounting. Results support previous literature on the discounting of delayed consequences by pathological gamblers, illustrate that the discounting of past rewards occurs in a similar fashion to the well-documented literature on the discounting of future rewards, and that magnitude of discounting can be altered.
PMCID: PMC2695330  PMID: 19949514
addiction; choice; delay discounting; pathological gambling; self-control
11.  An Efficient Operant Choice Procedure for Assessing Delay Discounting in Humans: Initial Validation in Cocaine-Dependent and Control Individuals 
Delay discounting is the decline in a consequence's control of behavior as a function of its delay, and may be a fundamental behavioral process in drug dependence. Human delay-discounting studies have usually relied on choices between hypothetical rewards. Some human tasks have assessed delay discounting using operant procedures with consequences provided during the task, as in nonhuman animal studies. However, these tasks have limitations such as long duration, potentially indeterminate data, or confounding the effect of delay with probability. A study in 20 cocaine-dependent volunteers and 20 demographically matched non-cocaine-dependent volunteers was designed to investigate a novel operant delay-discounting task providing monetary reinforcement by coin delivery throughout the task (Quick Discounting Operant Task; QDOT). Participants completed a hypothetical delay-discounting procedure, a potentially real reward delay-discounting procedure, and an existing operant delay-discounting task: the Experiential Discounting Task (EDT). The QDOT resulted in complete data for all participants, showed systematic effects of delay that were well described by a hyperbolic function, had a maximum duration of 17 min, and resulted in relatively little variability in session earnings. QDOT performance was significantly, positively correlated with performance on the EDT but not the other tasks. The QDOT resulted in an effect size between the groups that was similar to most other delay discounting tasks examined, and showed the cocaine-dependent participants to delay discount significantly more than the control participants. The QDOT is an efficient operant human delay-discounting task that may be useful in a variety of experimental settings.
PMCID: PMC3535463  PMID: 22329554
delay discounting; cocaine; real; hypothetical; human
12.  Regional White Matter Volumes Correlate with Delay Discounting 
PLoS ONE  2012;7(2):e32595.
A preference for immediate gratification is a central feature in addictive processes. However, the neural structures underlying reward delay tolerance are still unclear. Healthy participants (n = 121) completed a delay discounting questionnaire assessing the extent to which they prefer smaller immediate rewards to larger delayed reward after undergoing magnetic resonance imaging (MRI) scanning. Whole brain voxel-based morphometric analysis shows that delay discounting severity was negatively correlated with right prefrontal subgyral white matter volume and positively correlated with white matter volume in parahippocampus/hippocampus, after whole brain correction. This study might better our understanding of the neural basis of impulsivity and addiction.
PMCID: PMC3290657  PMID: 22393420
13.  Continuous theta burst stimulation of right dorsolateral prefrontal cortex induces changes in impulsivity level 
Brain stimulation  2009;3(3):170-176.
There is evidence that the right dorsolateral prefrontal cortex (DLPFC) may play a certain role in decision making related to reward value and time perception and, in particular, in the inhibitory control of impulsive decision making. Using the theta burst stimulation (TBS) and a delay discounting (DD) task, we investigated the potential role of right DLPFC in impulsive decision making defined by the rate of discounting delayed reward. Healthy right-handed volunteers underwent three stimulation sessions, intermittent TBS (iTBS), continuous TBS (cTBS), and sham. The steepness of the discount function (k-value), reaction time for choice and consistency were measured for each subjects. cTBS of the DLPFC reduced by 36.88 % the k-value of the DD task compared to sham condition. In contrast, iTBS did not affect impulsivity level. There were no changes neither in reaction time for choice nor consistency after either the iTBS or cTBS compared with the sham stimulation. These results demonstrate that cTBS-induced modulation of cortical excitability of the right DLPFC may affect and reduce impulsive decision making. These observations may provide some insights into the role of the right DLPFC in modulating impulsivity level and calculating reward value at different time scales under less ambiguous circumstances.
PMCID: PMC3707839  PMID: 20633446 CAMSID: cams3169
rTMS; theta burst stimulation; dorsolateral prefrontal cortex; decision making; impulsivity; delay discounting task
14.  Effect of continuous theta burst stimulation of the right dorsolateral prefrontal cortex on cerebral blood flow changes during decision making 
Brain stimulation  2012;5(2):116-123.
Decision making is a cognitive function relaying on a complex neural network. In particular, the right dorsolateral prefrontal cortex (DLPFC) plays a key role within this network. We used positron emission tomography (PET) combined with continuous theta burst transcranial magnetic stimulation (cTBS) to investigate neuronal and behavioral changes in normal volunteers while performing a delay discounting (DD) task. We aimed to test whether stimulation of right DLPFC would modify the activation pattern of the neural circuit underlying decision making during the DD task and influence discounting behavior.
We found that cTBS of the right DLPFC influenced decision making by reducing impulsivity and inducing participants to favor large but delayed rewards instead of immediate but small rewards. Stimulation also affected activation in several prefrontal areas associated with DD. In particular, we observed a reduced regional cerebral blood flow (rCBF) in the ipsilateral DLPFC (BA 46) extending into the rostral part of the prefrontal cortex (BA 10) as well as a disrupted relationship between impulsivity (k-value) and rCBF in these and other prefrontal areas.
These findings suggest that transcranial magnetic stimulation of the DLPFC influences the neural network underlying impulsive decision making behavior.
PMCID: PMC3707841  PMID: 22494829 CAMSID: cams3170
rTMS; Theta burst stimulation; Dorsolateral prefrontal cortex; Decision making; Impulsivity; Delay discounting task
15.  Parental inconsistency, impulsive choice and neural value representations in healthy adolescents 
Translational Psychiatry  2014;4(4):e382-.
A well-characterized potential marker for addiction is impulsive choice, stably measured by delay discounting (DD) paradigms. While genetic influences partly account for inter-individual variance in impulsivity, environmental factors such as parenting practices may have an important role. The present study investigates how inconsistent fulfillment of delayed reward promises impacts on DD. A combined correlational and experimental functional magnetic resonance imaging (fMRI) design was performed in a sample of 48 healthy adolescents (13–15 years). More specifically, neural activation during a DD task was investigated at two assessment points (T0 and T1). Adolescents' self-reports of parenting and substance use were assessed at T0. Between assessment points, we experimentally varied the reliability of delayed reward promises, measuring the impact of this intervention on DD and neural value processing at T1. In the correlational part, same-sex parent reward inconsistency was associated with steeper DD and an attenuated subjective value (SV) representation in the nucleus accumbens (NAcc) and ventromedial prefrontal cortex (vmPFC). Steeper DD was in turn associated with alcohol use during the past year. In the experimental part, the reward inconsistency manipulation resulted in an attenuation of the NAcc SV representation, similar to the parental inconsistency effect. Together, our correlational and experimental findings raise new light on how parents may influence their children's degree of impulsivity, making parenting a potential target in addiction prevention.
PMCID: PMC4012284  PMID: 24736798
16.  Neuroeconomics and Adolescent Substance Abuse: Individual Differences in Neural Networks and Delay Discounting 
Many adolescents with substance use problems show poor response to evidence based treatments. Treatment outcome has been associated with individual differences in impulsive decision making as reflected by delay discounting (DD) rates (preference for immediate rewards). Adolescents with higher rates of DD were expected to show greater neural activation in brain regions mediating impulsive/habitual behavioral choices and less activation in regions that mediate reflective/executive behavioral choices.
Thirty adolescents being treated for substance abuse completed a DD task optimized to balance choices of immediate versus delayed rewards and a control condition accounted for activation during magnitude valuation. A group independent component analysis on functional magnetic resonance imaging (fMRI) time courses identified neural networks engaged during DD. Network activity was correlated with individual differences in discounting rate.
Higher discounting rates were associated with diminished engagement of an executive attention control network involving the dorsolateral prefrontal cortex, dorsomedial prefrontal cortex, inferior parietal cortex, cingulate cortex, and precuneus. Higher discounting rates were also associated with less deactivation in a “bottom up” reward valuation network involving the amygdala, hippocampus, insula, and ventromedial prefrontal cortex. These 2 networks were significantly negatively correlated.
Results support relations between competing executive and reward valuation neural networks and temporal decision making, an important potentially modifiable risk factor relevant for prevention and treatment of adolescent substance abuse.
Clinical trial registration information—The Neuroeconomics of Behavioral Therapies for Adolescent Substance Abuse;; NCT01093898.
PMCID: PMC3712894  PMID: 23800488
adolescent substance abuse; delay discounting; functional magnetic resonance imaging (fMRI); neuroeconomics
17.  Biphasic Effects of Alcohol on Delay and Probability Discounting 
Delay discounting and probability discounting are behavioral economic indices of impulsive and risky decision making that have been associated with addictive behavior, but the acute biphasic effects of alcohol on these decision-making processes are not well understood. This study sought to investigate the biphasic effects of alcohol on delay and probability discounting across the ascending and descending limbs of the breath alcohol concentration (BAC) curve, which are respectively characterized by the stimulant and sedative effects of alcohol. Delay and probability discounting were measured at four time points (Baseline, Ascending, Descending, and Endpoint) across the BAC curve at two target alcohol doses (40 mg/dl and 80 mg/dl) in healthy adults (N = 23 and 27, for both doses, respectively). There was no significant effect of alcohol on delay discounting at either dose. Alcohol significantly affected probability discounting, such that reduced discounting for uncertain rewards was evident during the descending limb of the BAC curve at the lower dose (p<.05) and during both the ascending and descending limb of the BAC curve at the higher dose (p<.05). Thus, alcohol resulted in increased risky decision making, particularly during the descending limb which is primarily characterized by the sedative effects of alcohol. These findings suggest that the biphasic effects of alcohol across the ascending and descending limbs of the BAC have differential effects on behavior related to decision-making for probabilistic, but not delayed, rewards. Parallels to and distinctions from previous findings are discussed.
PMCID: PMC4050433  PMID: 23750692
delay discounting; probability discounting; impulsivity; alcohol use; behavioral economics
18.  Decisions for Others Become Less Impulsive the Further Away They Are on the Family Tree 
PLoS ONE  2012;7(11):e49479.
People tend to prefer a smaller immediate reward to a larger but delayed reward. Although this discounting of future rewards is often associated with impulsivity, it is not necessarily irrational. Instead it has been suggested that it reflects the decision maker’s greater interest in the ‘me now’ than the ‘me in 10 years’, such that the concern for our future self is about the same as for someone else who is close to us.
Methodology/Principal Findings
To investigate this we used a delay-discounting task to compare discount functions for choices that people would make for themselves against decisions that they think that other people should make, e.g. to accept $500 now or $1000 next week. The psychological distance of the hypothetical beneficiaries was manipulated in terms of the genetic coefficient of relatedness ranging from zero (e.g. a stranger, or unrelated close friend), .125 (e.g. a cousin), .25 (e.g. a nephew or niece), to .5 (parent or sibling).
The observed discount functions were steeper (i.e. more impulsive) for choices in which the decision-maker was the beneficiary than for all other beneficiaries. Impulsiveness of decisions declined systematically with the distance of the beneficiary from the decision-maker. The data are discussed with reference to the implusivity and interpersonal empathy gaps in decision-making.
PMCID: PMC3509068  PMID: 23209580
19.  Now or later? Striatum and insula activation to immediate versus delayed rewards 
Neuroimaging studies on delay discounting tasks that use reward delays ranging from minutes to days have implicated the insula and striatum in the processing of inter-temporal decisions. This study aimed at assessing whether these brain regions would also be involved in decision-making when subjects have to wait through the delays within the range of seconds. Employing functional magnetic resonance imaging (fMRI) in thirteen healthy volunteers, we repeatedly presented monetaryoptions with delays that differed within the range of multiple seconds. Using a region of interest approach, we found significant activation in the bilateral anterior insula and striatum when subjects chose either the immediate (smaller) or delayed (larger) option. In particular, insular activation was observed after the response and the delay, when the outcome of the immediate or the delayed choice was shown. Significantly greater activation was observed in the ventroanterior striatum while subjects chose the immediate, as opposed to the delayed, options, and also after receiving the outcome of waiting through the longer delay option. The evidence presented here indicates that both the ventral striatum and the insula are involved in the processing of choosing delay options as well as the consequences of choices with delays in the seconds’ range.
PMCID: PMC2907916  PMID: 20657814
delay discounting; decision making; time perception; reward; fMRI
20.  Impulsiveness as a timing disturbance: neurocognitive abnormalities in attention-deficit hyperactivity disorder during temporal processes and normalization with methylphenidate 
We argue that impulsiveness is characterized by compromised timing functions such as premature motor timing, decreased tolerance to delays, poor temporal foresight and steeper temporal discounting. A model illustration for the association between impulsiveness and timing deficits is the impulsiveness disorder of attention-deficit hyperactivity disorder (ADHD). Children with ADHD have deficits in timing processes of several temporal domains and the neural substrates of these compromised timing functions are strikingly similar to the neuropathology of ADHD. We review our published and present novel functional magnetic resonance imaging data to demonstrate that ADHD children show dysfunctions in key timing regions of prefrontal, cingulate, striatal and cerebellar location during temporal processes of several time domains including time discrimination of milliseconds, motor timing to seconds and temporal discounting of longer time intervals. Given that impulsiveness, timing abnormalities and more specifically ADHD have been related to dopamine dysregulation, we tested for and demonstrated a normalization effect of all brain dysfunctions in ADHD children during time discrimination with the dopamine agonist and treatment of choice, methylphenidate. This review together with the new empirical findings demonstrates that neurocognitive dysfunctions in temporal processes are crucial to the impulsiveness disorder of ADHD and provides first evidence for normalization with a dopamine reuptake inhibitor.
PMCID: PMC2685816  PMID: 19487194
impulsiveness; timing; time perception; temporal discounting; attention-deficit hyperactivity disorder; methylphenidate
21.  Isolating the delay component of impulsive choice in adolescent rats 
Impulsive choice—the preference for small immediate rewards over larger delayed rewards—has been linked to various psychological conditions ranging from behavioral disorders to addiction. These links highlight the critical need to dissect the various components of this multifaceted behavioral trait. Delay discounting tasks allow researchers to study an important factor of this behavior: how the subjective value of a rewards changes over a delay period. However, existing methods of delay discounting include a confound of different reward sizes within the procedure. Here we present a new approach of using a single constant reward size to assess delay discounting. A complementary approach could hold delay constant and assess the utility of changing quantities of a reward. Isolating these behavioral components can advance our ability to explore the behavioral complexity of impulsive choice. We present in detail the methods for isolating delay, and further capitalize on this method by pairing it with a standard peak interval task to test whether individual variation in delay discounting can be explained by differences in perception of time in male and female adolescent rats. We find that rats that were more precise in discriminating time intervals were also less impulsive in their choice. Our data suggest that differences in timing and delay discounting are not causally related, but instead are more likely influenced by a common factor. Further, the mean-level change in our measure between post-natal day 28 and 42 suggests this test may be capturing a developmental change in this factor. In summary, this new method of isolating individual components of impulsive choice (delay or quantity) can be efficiently applied in either adolescent or adult animal models and may help elucidate the mechanisms underlying impulsivity and its links to psychological disorders.
PMCID: PMC3902300  PMID: 24478644
impulsive choice; delay discounting; peak interval; adolescence; sex differences
22.  Notes on Discounting 
In general, if a variable can be expressed as a function of its own maximum value, that function may be called a discount function. Delay discounting and probability discounting are commonly studied in psychology, but memory, matching, and economic utility also may be viewed as discounting processes. When they are so viewed, the discount function obtained is hyperbolic in form. In some cases the effective discounting variable is proportional to the physical variable on which it is based. For example, in delay discounting, the physical variable, delay (D), may enter into the hyperbolic equation as kD. In many cases, however, the discounting data are not well described with a single-parameter discount function. A much better fit is obtained when the effective variable is a power function of the physical variable (kDs in the case of delay discounting). This power-function form fits the data of delay, probability, and memory discounting as well as other two-parameter discount functions and is consistent with both the generalized matching law and maximization of a constant-elasticity-of-substitution utility function.
PMCID: PMC1459845  PMID: 16776060
discounting; delay discounting; memory discounting; social discounting; probability discounting; matching; rational behavior; utility maximization
23.  Effort discounting in human nucleus accumbens 
A great deal of behavioral and economic research suggests that the value attached to a reward stands in inverse relation to the amount of effort required to obtain it, a principle known as effort discounting. In the current report, we present the first direct evidence for a neural analogue of effort discounting. Functional magnetic resonance imaging was used to measure neural responses to monetary rewards in the human nucleus accumbens, a structure previously demonstrated to encode reference-dependent reward information. The magnitude of accumbens activation was found to vary with reward outcome, but also with the degree of mental effort demanded to obtain individual rewards. For a fixed level of reward, accumbens was less strongly activated following a high demand for effort than following a lower demand. The magnitude of this effect was noted to correlate with preceding activation in the dorsal anterior cingulate cortex, a region that has been proposed to monitor information-processing demands and to mediate in the subjective experience of effort.
PMCID: PMC2744387  PMID: 19246324
24.  Impact of glutamate levels on neuronal response and cognitive abilities in schizophrenia 
NeuroImage : Clinical  2014;4:576-584.
Schizophrenia is characterized by impaired cognitive functioning, and brain regions involved in cognitive control processes show marked glutamatergic abnormalities. However, it is presently unclear whether aberrant neuronal response is directly related to the observed deficits at the metabolite level in schizophrenia. Here, 17 medicated schizophrenia patients and 17 matched healthy participants underwent functional magnetic resonance imaging (fMRI) when performing an auditory cognitive control task, as well as proton magnetic resonance spectroscopy (1H-MRS) in order to assess resting-state glutamate in the anterior cingulate cortex. The combined fMRI–1H-MRS analysis revealed that glutamate differentially predicted cortical blood-oxygen level-dependent (BOLD) response in patients and controls. While we found a positive correlation between glutamate and BOLD response bilaterally in the inferior parietal lobes in the patients, the corresponding correlation was negative in the healthy control participants. Further, glutamate levels predicted task performance in patients, such that lower glutamate levels were related to impaired cognitive control functioning. This was not seen for the healthy controls. These findings suggest that schizophrenia patients have a glutamate-related dysregulation of the brain network supporting cognitive control functioning. This could be targeted in future research on glutamatergic treatment of cognitive symptoms in schizophrenia.
•Neuronal processing of cognitive control is different in schizophrenia patients (SZ).•Cingulum glutamate levels predict the degree of parietal neuronal response.•Lower glutamate predicts poorer cognitive control abilities in SZ.•SZ have a glutamate-related neuronal dysregulation of cognitive control processing.
PMCID: PMC3989526  PMID: 24749064
Combined fMRI–MRS; 1H-MRS; Cognitive control; Anterior cingulate cortex; Inferior parietal lobe; Glutamate; BOLD; Connectivity
25.  Prefrontal cortical–striatal dopamine receptor mRNA expression predicts distinct forms of impulsivity 
The European journal of neuroscience  2013;37(11):1779-1788.
Variation in dopamine receptor levels has been associated with different facets of impulsivity. To further delineate the neural substrates underlying impulsive action (inability to withhold a prepotent motor response) and impulsive choice (delay aversion), we characterised rats in the Differential Reinforcement of Low Rates of Responding task and a delay discounting task. We also measured performance on an effort-based discounting task. We then assessed D1 and D2 dopamine receptor mRNA expression in subregions of the prefrontal cortex and nucleus accumbens using in situ hybridisation, and compared these data with behavioral performance. Expression of D1 and D2 receptor mRNA in distinct brain regions was predictive of impulsive action. A dissociation within the nucleus accumbens was observed between subregions and receptor subtypes; higher D1 mRNA expression in the shell predicted greater impulsive action, whereas lower D2 mRNA expression in the core predicted greater impulsive action. We also observed a negative correlation between impulsive action and D2 mRNA expression in the prelimbic cortex. Interestingly, a similar relationship was present between impulsive choice and prelimbic cortex D2 mRNA, despite the fact that behavioral indices of impulsive action and impulsive choice were uncorrelated. Finally, we found that both high D1 mRNA expression in the insular cortex and low D2 mRNA expression in the infralimbic cortex were associated with willingness to exert effort for rewards. Notably, dopamine receptor mRNA in these regions was not associated with either facet of impulsivity. The data presented here provide novel molecular and neuroanatomical distinctions between different forms of impulsivity, as well as effort-based decision-making.
PMCID: PMC3973541  PMID: 23510331
delay discounting; effort discounting; impulsive action; impulsive choice; rat

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