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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.
doi:10.1016/j.neuroimage.2008.11.004
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 
Background
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.
Methods
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.
Results
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.
Conclusions
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.
doi:10.1111/j.1530-0277.2011.01455.x
PMCID: PMC3117198  PMID: 21676001
Delay discounting; intertemporal choice; impulsivity; delayed gratification
3.  Frontal Hyperconnectivity Related to Discounting and Reversal Learning in Cocaine Subjects 
Biological psychiatry  2011;69(11):1117-1123.
BACKGROUND
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.
METHODS
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.
RESULTS
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.
CONCLUSIONS
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.
doi:10.1016/j.biopsych.2011.01.008
PMCID: PMC3090521  PMID: 21371689
cocaine; functional connectivity; anterior cingulate; delayed discount; reversal learning; frontal
4.  Cortical activation during delay discounting in abstinent methamphetamine dependent individuals 
Psychopharmacology  2008;201(2):183-193.
Background
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.
Methods
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.
Results
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.
Conclusions
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.
doi:10.1007/s00213-008-1261-1
PMCID: PMC2835463  PMID: 18685833
Methamphetamine; Delay discounting; Brain imaging
5.  ALTERING THE MAGNITUDE OF DELAY DISCOUNTING BY PATHOLOGICAL GAMBLERS 
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.
doi:10.1901/jaba.2009.42-269
PMCID: PMC2695330  PMID: 19949514
addiction; choice; delay discounting; pathological gambling; self-control
6.  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.
doi:10.1371/journal.pone.0032595
PMCID: PMC3290657  PMID: 22393420
7.  Decisions for Others Become Less Impulsive the Further Away They Are on the Family Tree 
PLoS ONE  2012;7(11):e49479.
Background
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).
Conclusions/Significance
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.
doi:10.1371/journal.pone.0049479
PMCID: PMC3509068  PMID: 23209580
8.  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.
doi:10.1037/a0017252
PMCID: PMC2907916  PMID: 20657814
delay discounting; decision making; time perception; reward; fMRI
9.  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.
doi:10.1098/rstb.2009.0014
PMCID: PMC2685816  PMID: 19487194
impulsiveness; timing; time perception; temporal discounting; attention-deficit hyperactivity disorder; methylphenidate
10.  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.
doi:10.1901/jeab.2006.85-05
PMCID: PMC1459845  PMID: 16776060
discounting; delay discounting; memory discounting; social discounting; probability discounting; matching; rational behavior; utility maximization
11.  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.
doi:10.3758/CABN.9.1.16
PMCID: PMC2744387  PMID: 19246324
12.  Delay Discounting and Future-directed Thinking in Anhedonic Individuals 
Anhedonia (lack of reactivity to pleasurable stimuli) and a negatively skewed view of the future are important components of depression that could affect economic decisions in depressed individuals. Delay discounting paradigms might be useful for probing putative affective and cognitive underpinnings of such decisions. As a first step to evaluate whether difficulties experiencing pleasure might affect delay discounting, 36 undergraduate students with varying levels of anhedonia performed a delay discounting task in which they made choices between a small immediate and larger future monetary reward. Increasing levels of anhedonia (Snaith-Hamilton Pleasure Scale) were negatively associated with delay discounting rate, indicating that anhedonic individuals tended to choose the larger, albeit delayed reward. These correlations remained after controlling for variables previously linked to delay discounting (working memory capacity and impulsivity) and pessimistic future-directed thinking. The current findings provide preliminary evidence indicating that anhedonic individuals make less myopic decisions about their future, possibly due to their decreased responsiveness to immediate rewards.
doi:10.1016/j.jbtep.2010.02.003
PMCID: PMC2862767  PMID: 20219184
Delay discounting; Anhedonia; Impulsivity; Working memory; Depression; Future-directed thinking
13.  Does delay discounting play an etiological role in smoking or is it a consequence of smoking? 
Drug and alcohol dependence  2009;103(3):99-106.
Although higher delay discounting rates have been linked to cigarette smoking, little is known about the stability of delay discounting, whether delay discounting promotes smoking acquisition, whether smoking contributes to impulsive choices, or if different relationships exist in distinct subgroups. This study sought to fill these gaps within a prospective longitudinal cohort study (N=947) spanning mid adolescence to young adulthood (age 15 to 21 years old). Smoking and delay discounting were measured across time. Covariates included peer and household smoking, academic performance, depression, novelty seeking, inattention and hyperactivity/impulsivity symptoms, and alcohol and marijuana use. The associated processes Latent Growth Curve Modeling (LGCM) with paths from the delay discounting level factor (baseline measure) and the trend factor (slope) to the smoking trend factor (slope) fit the data well, X2(19, n=947) = 15.37, p=.70, CFI=1.00, RMSEA=0, WRMR=.36. The results revealed that delay discounting did not change significantly across time. Baseline delay discounting had a significant positive effect on smoking trend (β=.08, z=2.16, p=.03). A standard deviation (SD=1.41) increase in baseline delay discounting resulted in an 11% increase (OR=1.11, 95% CI= 1.03, 1.23) in the odds of smoking uptake. The alternative path LCGM revealed that smoking did not significantly impact delay discounting (p’s > .05). Growth Mixture Modeling identified three smoking trajectories: nonsmokers, early/fast smoking adopters, and slow smoking progressors. Delay discounting was higher in the smoking versus nonsmoking trajectories, but did not discriminate between the smoking trajectories, despite different acquisition patterns. Delay discounting may provide a variable by which to screen for smoking vulnerability and help identify subgroups to target for more intensive smoking prevention efforts that include novel behavioral components directed toward aspects of impulsivity.
doi:10.1016/j.drugalcdep.2008.12.019
PMCID: PMC2743449  PMID: 19443136
Delay discounting; smoking acquisition
14.  Individual Differences in Delay Discounting Under Acute Stress: The Role of Trait Perceived Stress 
Delay discounting refers to the reduction of the value of a future reward as the delay to that reward increases. The rate at which individuals discount future rewards varies as a function of both individual and contextual differences, and high delay discounting rates have been linked with problematic behaviors, including drug abuse and gambling. The current study investigated the effects of acute anticipatory stress on delay discounting, while considering two important factors: individual perceptions of stress and whether the stressful situation is future-focused or present-focused. Half of the participants experienced acute stress by anticipating giving a videotaped speech. This stress was either future-oriented (speech about future job) or present-oriented (speech about physical appearance). They then performed a delay discounting task, in which they chose between smaller, immediate rewards, and larger, delayed rewards. Their scores on the Perceived Stress Scale were also collected. The way in which one appraises stressful situations interacts with acute stress to influence choices; under stressful conditions, delay discounting rate was highest in individuals with low trait perceived stress and lowest for individuals with high trait perceived stress. This result might be related to individual variation in reward responsiveness under stress. Furthermore, the time orientation of the task interacted with its stressfulness to affect the individual’s propensity to choose immediate rewards. These findings add to our understanding of the intermediary factors between stress and decision-making.
doi:10.3389/fpsyg.2012.00251
PMCID: PMC3400439  PMID: 22833731
delay discounting; stress; decision-making; future orientation; perceived stress
15.  Humans Can Adopt Optimal Discounting Strategy under Real-Time Constraints 
PLoS Computational Biology  2006;2(11):e152.
Critical to our many daily choices between larger delayed rewards, and smaller more immediate rewards, are the shape and the steepness of the function that discounts rewards with time. Although research in artificial intelligence favors exponential discounting in uncertain environments, studies with humans and animals have consistently shown hyperbolic discounting. We investigated how humans perform in a reward decision task with temporal constraints, in which each choice affects the time remaining for later trials, and in which the delays vary at each trial. We demonstrated that most of our subjects adopted exponential discounting in this experiment. Further, we confirmed analytically that exponential discounting, with a decay rate comparable to that used by our subjects, maximized the total reward gain in our task. Our results suggest that the particular shape and steepness of temporal discounting is determined by the task that the subject is facing, and question the notion of hyperbolic reward discounting as a universal principle.
Synopsis
When we make a choice between two options, we compare the values of their outcomes and select the option with a larger value. However, what if one option leads to a larger delayed reward and the other leads to a smaller more immediate reward? Naturally, we assign a larger value for a larger reward, but it is “discounted” if the reward is to be delivered later. Thus, the value is a monotonically decreasing function of the delays. Previous behavioral studies have repeatedly demonstrated that humans and animals discount delayed rewards hyperbolically. This has practical importance, as hyperbolic discounting can sometimes lead to “irrational” preference reversal: for instance, an individual may prefer two apples in 51 days to one apple in 50 days, but if the days come closer, he prefers one apple today to two apples tomorrow. On the contrary, exponential discounting is always “rational,” as it predicts constant preference. Here, in a new task that mimics animal foraging, and that uses delayed monetary rewards, Schweighofer and colleagues showed that humans can also discount reward exponentially. Furthermore, it is remarkable that by adopting exponential discounting, their subjects maximized their total gain. Thus, depending on the task at hand, the authors' study suggests that humans can flexibly choose the type of reward discounting, and can exhibit rational behavior that maximizes long-term gains.
doi:10.1371/journal.pcbi.0020152
PMCID: PMC1635539  PMID: 17096592
16.  Behavioral and Neuroeconomics of Drug Addiction: Competing Neural Systems and Temporal Discounting Processes 
Drug and alcohol dependence  2006;90(Suppl 1):S85-S91.
We review behavioral- and neuroeconomic research that identifies temporal discounting as an important component in the development and maintenance of drug addiction. First we review behavioral economic research that explains and documents the contribution of temporal discounting to addiction. This is followed with recent insights from neuroeconomics that may provide an explanation of why drug dependent individuals discount the future. Specifically, neuroeconomics has identified two competing neural systems that are related to temporal discounting using brain-imaging techniques that examine the relative activation of different brain regions for temporal discounting. According to the competing neural systems account, choices for delayed outcomes are related to the prefrontal cortex (i.e., the “executive system”) and choices for immediate outcomes are related to the limbic brain regions (i.e., the “impulsive system”). Temporal discounting provides a useful framework for future imaging research, and suggests a novel approach to designing effective drug dependence prevention and treatment programs.
doi:10.1016/j.drugalcdep.2006.09.016
PMCID: PMC2033431  PMID: 17101239
Behavioral economics; Neuroeconomics; Temporal discounting; Functional magnetic resonance imaging (fMRI); Addiction; Decision making
17.  Short-Term Temporal Discounting of Reward Value in Human Ventral Striatum 
Journal of Neurophysiology  2009;101(3):1507-1523.
Delayed rewards lose their value for economic decisions and constitute weaker reinforcers for learning. Temporal discounting of reward value already occurs within a few seconds in animals, which allows investigations of the underlying neurophysiological mechanisms. However, it is difficult to relate these mechanisms to human discounting behavior, which is usually studied over days and months and may engage different brain processes. Our study aimed to bridge the gap by using very short delays and measuring human functional magnetic resonance responses in one of the key reward centers of the brain, the ventral striatum. We used psychometric methods to assess subjective timing and valuation of monetary rewards with delays of 4.0–13.5 s. We demonstrated hyperbolic and exponential decreases of striatal responses to reward predicting stimuli within this time range, irrespective of changes in reward rate. Lower reward magnitudes induced steeper behavioral and striatal discounting. By contrast, striatal responses following the delivery of reward reflected the uncertainty in subjective timing associated with delayed rewards rather than value discounting. These data suggest that delays of a few seconds affect the neural processing of predicted reward value in the ventral striatum and engage the temporal sensitivity of reward responses. Comparisons with electrophysiological animal data suggest that ventral striatal reward discounting may involve dopaminergic and orbitofrontal inputs.
doi:10.1152/jn.90730.2008
PMCID: PMC2666398  PMID: 19164109
18.  Domain independence and stability in young and older adults' discounting of delayed rewards 
Behavioural processes  2011;87(3):253-259.
Individual discounting rates for different types of delayed reward are typically assumed to reflect a single, underlying trait of impulsivity. Recently, we showed that discounting rates are orders of magnitude steeper for directly consumable liquid rewards than for monetary rewards (Jimura et al. 2009), raising the question of whether discounting rates for different types of reward covary at the individual level. Accordingly, the present study examined the relation between discounting of hypothetical money and real liquid rewards in young adults (Experiment 1) and older adults (Experiment 2). At the group level, young adults discounted monetary rewards more steeply than the older adults, but the reverse pattern was observed with liquid rewards. At the individual level, the rates at which young and older participants discounted each reward type were stable over a two- to fifteen-week interval (rs >.70), but there was no significant correlation between the rates at which they discounted the two reward types. These results suggest that although similar decision-making processes may underlie the discounting of different types of rewards, the rates at which individuals discount money and directly consumable rewards may reflect separate, stable traits, rather than a single trait of impulsivity.
doi:10.1016/j.beproc.2011.04.006
PMCID: PMC3138910  PMID: 21550384
delay discounting; domain independence; stability; monetary reward; consumable reward; aging
19.  Three Systems of Insular Functional Connectivity Identified with Cluster Analysis 
Cerebral Cortex (New York, NY)  2010;21(7):1498-1506.
Despite much research on the function of the insular cortex, few studies have investigated functional subdivisions of the insula in humans. The present study used resting-state functional connectivity magnetic resonance imaging (MRI) to parcellate the human insular lobe based on clustering of functional connectivity patterns. Connectivity maps were computed for each voxel in the insula based on resting-state functional MRI (fMRI) data and segregated using cluster analysis. We identified 3 insular subregions with distinct patterns of connectivity: a posterior region, functionally connected with primary and secondary somatomotor cortices; a dorsal anterior to middle region, connected with dorsal anterior cingulate cortex, along with other regions of a previously described control network; and a ventral anterior region, primarily connected with pregenual anterior cingulate cortex. Applying these regions to a separate task data set, we found that dorsal and ventral anterior insula responded selectively to disgusting images, while posterior insula did not. These results demonstrate that clustering of connectivity patterns can be used to subdivide cerebral cortex into anatomically and functionally meaningful subregions; the insular regions identified here should be useful in future investigations on the function of the insula.
doi:10.1093/cercor/bhq186
PMCID: PMC3116731  PMID: 21097516
disgust; insula; parcellation; resting state
20.  An Argument Against Dual Valuation System Competition: Cognitive Capacities Supporting Future Orientation Mediate Rather Than Compete With Visceral Motivations 
The dynamic inconsistency of preference is well documented in behavioral research, but its basis remains controversial. In this article, we summarize recent functional magnetic resonance imaging (fMRI) work in the domain of intertemporal choice, specifically considering evidence bearing on the hypothesis that delay discounting in humans is determined by competition between an evolutionarily older system that discounts precipitously with delay (System 1), and a newer system that exhibits very little discounting (System 2). We argue that neuroimaging evidence does not support the hypothesized separate and competing value systems. While it is clear that the sophisticated cognitive capacities that lead to greater valuation of larger later alternatives (e.g., selective attention and self-signaling) depend critically on neocortical structures, these capacities affect intertemporal choice through mediation of (rather than competition with) older cortical and subcortical structures central to reward and motivation. Taken together, neuroimaging evidence supports the alternative hypothesis that intertemporal choice is guided by a single valuation system.
doi:10.1037/a0016827
PMCID: PMC3169839  PMID: 21909453
self-control; delay discounting; self-signaling; neuroeconomics
21.  Delay Discounting as a Mediator of the Relationship between Perceived Stress and Cigarette Smoking Status in Adolescents 
Behavioural pharmacology  2009;20(5-6):455-460.
There has been a wealth of research providing evidence for the relationship between stress and cigarette smoking during adolescence. Despite this knowledge, little is known about possible behavioral mechanisms by which stress exerts its influence on the decision to smoke. This study sought to examine one such behavioral characteristic, delay discounting, that may mediate the relationship between stress and cigarette smoking. Delay discounting generally refers to the discounting of value for outcomes because they are delayed; and high rates of delay discounting have been linked to impulsive behavior. For the current research, adolescent smokers (n = 50) and nonsmokers (n = 50) were compared using a self-report measure of perceived stress and a laboratory assessment of delay discounting. Smokers tended to report higher levels of stress and to discount more by delay, and there was a significant association between reported stress and delay discounting. Additionally, delay discounting mediated the relationship between stress and cigarette smoking status. These results suggest that discounting by delay may be a behavior through which stress exerts influence on an adolescent’s decision to smoke.
doi:10.1097/FBP.0b013e328330dcff
PMCID: PMC3209706  PMID: 19730366
perceived stress; delay discounting; cigarette smoking; adolescents
22.  A hyperbolic decay of subjective probability of obtaining delayed rewards 
Background
Hyperbolic discounting of delayed and probabilistic outcomes has drawn attention in psychopharmacology and neuroeconomics. Sozou's evolutionary theory proposed that hyperbolic delay discounting may be totally attributable to aversion to a decrease in subjective probability of obtaining delayed rewards (SP) which follows a hyperbolic decay function. However, to date, no empirical study examined the hypothesis, although this investigation is important for elucidating the roles of impatience, precaution, and uncertainty aversion in delay discounting processes.
Methods
In order to (i) determine the functional form of the relation between delay until receipt and SP, and (ii) examine whether delay discounting is attributable to a decrease in SP, we assessed the subjects' SP and their delay and probability discounting. We examined the fitness of hyperbolic and exponential functions to the assessed SP, and relations between the SP, and delay/probability discounting, and subjective-probability discounting for delayed rewards.
Results
The results demonstrated (a) SP decayed hyperbolically as delay increases, (b) a decay of SP was associated with delay discounting, and (c) subjective-probability discounting did not significantly correlate with delay discounting.
Conclusion
Our results demonstrated (i) hyperbolic decay of SP is related to delay discounting, and (ii) delay discounting is, however, not attributable to precautious foresight in intertemporal choice. Further, a novel parameter of pure time preference is proposed.
doi:10.1186/1744-9081-3-52
PMCID: PMC2072955  PMID: 17892596
23.  Delay discounting: Trait variable? 
Behavioural Processes  2011;87(1):1-9.
Delay discounting refers to the tendency for outcomes that are remote in time to have less value than more immediate outcomes. Steep discounting of delayed outcomes is associated with a variety of social maladies. The degree of sensitivity to delayed outcomes may be a stable and pervasive individual characteristic. In analyses of archival data, the present study found positive correlations between the degree of delay discounting for one outcome (as measured by the Area Under the Curve), and the degree of discounting for other outcomes. Along with additional evidence reviewed, these data suggest that delay discounting may be considered a personality trait. Recent research in epigenetics, neuroscience, and behavior suggests delay discounting may prove to be a beneficial target for therapeutic attempts to produce global reductions in impulsivity related to delay discounting.
doi:10.1016/j.beproc.2011.02.007
PMCID: PMC3266171  PMID: 21385637
delay discounting; impulsivity; humans; personality; trait
24.  Default network connectivity reflects the level of consciousness in non-communicative brain-damaged patients 
Brain  2009;133(1):161-171.
The ‘default network’ is defined as a set of areas, encompassing posterior-cingulate/precuneus, anterior cingulate/mesiofrontal cortex and temporo-parietal junctions, that show more activity at rest than during attention-demanding tasks. Recent studies have shown that it is possible to reliably identify this network in the absence of any task, by resting state functional magnetic resonance imaging connectivity analyses in healthy volunteers. However, the functional significance of these spontaneous brain activity fluctuations remains unclear. The aim of this study was to test if the integrity of this resting-state connectivity pattern in the default network would differ in different pathological alterations of consciousness. Fourteen non-communicative brain-damaged patients and 14 healthy controls participated in the study. Connectivity was investigated using probabilistic independent component analysis, and an automated template-matching component selection approach. Connectivity in all default network areas was found to be negatively correlated with the degree of clinical consciousness impairment, ranging from healthy controls and locked-in syndrome to minimally conscious, vegetative then coma patients. Furthermore, precuneus connectivity was found to be significantly stronger in minimally conscious patients as compared with unconscious patients. Locked-in syndrome patient’s default network connectivity was not significantly different from controls. Our results show that default network connectivity is decreased in severely brain-damaged patients, in proportion to their degree of consciousness impairment. Future prospective studies in a larger patient population are needed in order to evaluate the prognostic value of the presented methodology.
doi:10.1093/brain/awp313
PMCID: PMC2801329  PMID: 20034928
Default mode; fMRI; coma; vegetative state; minimally conscious state
25.  Delay Discounting of Cocaine by Rhesus Monkeys 
The present, subjective value of a reinforcer typically decreases as a function of the delay to its receipt, a phenomenon termed delay discounting. Delay discounting, which is assumed to reflect impulsivity, is hypothesized to play an important role in drug abuse. The present study examined delay discounting of cocaine injections by rhesus monkeys. Subjects were studied on a discrete-trials task in which they chose between 2 doses of cocaine: a smaller, immediate dose and a larger, delayed dose. The immediate dose varied between 0.012 and 0.4 mg/kg/injection, whereas the delayed dose was always 0.2 mg/kg/injection and was delivered after a delay that varied between 0 and 300 s in different conditions. At each delay, the point at which a monkey chose the immediate and delayed doses equally often (i.e., the ED50) provided a measure of the present, subjective value of the delayed dose. Dose–response functions for the immediate dose shifted to the left as delay increased. The amount of the immediate dose predicted to be equal in subjective value to the delayed dose decreased as a function of the delay, and hyperbolic discounting functions provided good fits to the data (median R2 = .86). The current approach may provide the basis for an animal model of the effect of delay on the subjective value of drugs of abuse.
doi:10.1037/1064-1297.15.3.238
PMCID: PMC3107575  PMID: 17563210
delay discounting; cocaine; impulsivity; choice; monkeys

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