The Role of Serotonin in the Regulation of Patience and Impulsivity

Classic theories suggest that central serotonergic neurons are involved in the behavioral inhibition that is associated with the prediction of negative rewards or punishment. Failed behavioral inhibition can cause impulsive behaviors. However, the behavioral inhibition that results from predicting punishment is not sufficient to explain some forms of impulsive behavior. In this article, we propose that the forebrain serotonergic system is involved in “waiting to avoid punishment” for future punishments and “waiting to obtain reward” for future rewards. Recently, we have found that serotonergic neurons increase their tonic firing rate when rats await food and water rewards and conditioned reinforcer tones. The rate of tonic firing during the delay period was significantly higher when rats were waiting for rewards than for tones, and rats were unable to wait as long for tones as for rewards. These results suggest that increased serotonergic neuronal firing facilitates waiting behavior when there is the prospect of a forthcoming reward and that serotonergic activation contributes to the patience that allows rats to wait longer. We propose a working hypothesis to explain how the serotonergic system regulates patience while waiting for future rewards.     

Prospective Optimization with Limited Resources

The future is uncertain because some forthcoming events are unpredictable and also because our ability to foresee the myriad consequences of our own actions is limited. Here we studied how humans select actions under such extrinsic and intrinsic uncertainty, in view of an exponentially expanding number of prospects on a branching multivalued visual stimulus. A triangular grid of disks of different sizes scrolled down a touchscreen at a variable speed. The larger disks represented larger rewards. The task was to maximize the cumulative reward by touching one disk at a time in a rapid sequence, forming an upward path across the grid, while every step along the path constrained the part of the grid accessible in the future. This task captured some of the complexity of natural behavior in the risky and dynamic world, where ongoing decisions alter the landscape of future rewards. By comparing human behavior with behavior of ideal actors, we identified the strategies used by humans in terms of how far into the future they looked (their “depth of computation”) and how often they attempted to incorporate new information about the future rewards (their “recalculation period”). We found that, for a given task difficulty, humans traded off their depth of computation for the recalculation period. The form of this tradeoff was consistent with a complete, brute-force exploration of all possible paths up to a resource-limited finite depth. A step-by-step analysis of the human behavior revealed that participants took into account very fine distinctions between the future rewards and that they abstained from some simple heuristics in assessment of the alternative paths, such as seeking only the largest disks or avoiding the smaller disks. The participants preferred to reduce their depth of computation or increase the recalculation period rather than sacrifice the precision of computation.     

The Value of Emotion: How Does Episodic Prospection Modulate Delay Discounting?

Background

Humans often show impatience when making intertemporal choice for monetary rewards, preferring small rewards delivered immediately to larger rewards delivered after a delay, which reflects a fundamental psychological principle: delay discounting. However, we propose that episodic prospection humans can vividly envisage exerts a strong and broad influence on individuals'' delay discounting. Specifically, episodic prospection may affect individuals'' intertemporal choice by the negative or positive emotion of prospection.

Methodology/Principal Findings

The present study explored how episodic prospection modulated delay discounting by emotion. Study 1 showed that participants were more inclined to choose the delayed but larger rewards when they imaged positive future events than when they did not image events; Study 2 showed that participants were more inclined to choose the immediate but smaller rewards when they imaged negative future events than when they did not image events; In contrast, study 3 showed that choice preferences of participants when they imaged neutral future events were the same as when they did not image events.

Conclusions/Significance

By manipulating the emotion valence of episodic prospection, our findings suggested that positive emotion made individuals tend to choose delayed rewards, while negative emotion made individuals tend to choose immediate rewards. Only imaging events with neutral emotion did not affect individuals'' choice preference. Thus, the valence of imaged future events'' emotion might play an important role in individuals'' intertemporal choice. It is possible that the valence of emotion may affect the changed direction (promote or inhibit) of individuals'' delay discounting, while the ability to image future events affects the changed degree of individuals'' delay discounting.     

Neural Sensitivity to Absolute and Relative Anticipated Reward in Adolescents

Adolescence is associated with a dramatic increase in risky and impulsive behaviors that have been attributed to developmental differences in neural processing of rewards. In the present study, we sought to identify age differences in anticipation of absolute and relative rewards. To do so, we modified a commonly used monetary incentive delay (MID) task in order to examine brain activity to relative anticipated reward value (neural sensitivity to the value of a reward as a function of other available rewards). This design also made it possible to examine developmental differences in brain activation to absolute anticipated reward magnitude (the degree to which neural activity increases with increasing reward magnitude). While undergoing fMRI, 18 adolescents and 18 adult participants were presented with cues associated with different reward magnitudes. After the cue, participants responded to a target to win money on that trial. Presentation of cues was blocked such that two reward cues associated with $.20, $1.00, or $5.00 were in play on a given block. Thus, the relative value of the $1.00 reward varied depending on whether it was paired with a smaller or larger reward. Reflecting age differences in neural responses to relative anticipated reward (i.e., reference dependent processing), adults, but not adolescents, demonstrated greater activity to a $1 reward when it was the larger of the two available rewards. Adults also demonstrated a more linear increase in ventral striatal activity as a function of increasing absolute reward magnitude compared to adolescents. Additionally, reduced ventral striatal sensitivity to absolute anticipated reward (i.e., the difference in activity to medium versus small rewards) correlated with higher levels of trait Impulsivity. Thus, ventral striatal activity in anticipation of absolute and relative rewards develops with age. Absolute reward processing is also linked to individual differences in Impulsivity.     

Temporal discounting predicts risk sensitivity in rhesus macaques

Humans and animals tend both to avoid uncertainty and to prefer immediate over future rewards. The comorbidity of psychiatric disorders such as impulsivity, problem gambling, and addiction suggests that a common mechanism may underlie risk sensitivity and temporal discounting. Nonetheless, the precise relationship between these two traits remains largely unknown. To examine whether risk sensitivity and temporal discounting reflect a common process, we recorded choices made by two rhesus macaques in a visual gambling task while we varied the delay between trials. We found that preference for the risky option declined with increasing delay between sequential choices in the task, even when all other task parameters were held constant. These results were quantitatively predicted by a model that assumed that the subjective expected utility of the risky option is evaluated based on the expected time of the larger payoff. The importance of the larger payoff in this model suggests that the salience of larger payoffs played a critical role in determining the value of risky options. These data suggest that risk sensitivity may be a product of other cognitive processes, and specifically that myopia for the future and the salience of jackpots control the propensity to take a gamble.     

Mentalizing deficits constrain belief in a personal God

Religious believers intuitively conceptualize deities as intentional agents with mental states who anticipate and respond to human beliefs, desires and concerns. It follows that mentalizing deficits, associated with the autistic spectrum and also commonly found in men more than in women, may undermine this intuitive support and reduce belief in a personal God. Autistic adolescents expressed less belief in God than did matched neuro-typical controls (Study 1). In a Canadian student sample (Study 2), and two American national samples that controlled for demographic characteristics and other correlates of autism and religiosity (Study 3 and 4), the autism spectrum predicted reduced belief in God, and mentalizing mediated this relationship. Systemizing (Studies 2 and 3) and two personality dimensions related to religious belief, Conscientiousness and Agreeableness (Study 3), failed as mediators. Mentalizing also explained the robust and well-known, but theoretically debated, gender gap in religious belief wherein men show reduced religious belief (Studies 2-4).     

Prefrontal coding of temporally discounted values during intertemporal choice

Reward from a particular action is seldom immediate, and the influence of such delayed outcome on choice decreases with delay. It has been postulated that when faced with immediate and delayed rewards, decision makers choose the option with maximum temporally discounted value. We examined the preference of monkeys for delayed reward in an intertemporal choice task and the neural basis for real-time computation of temporally discounted values in the dorsolateral prefrontal cortex. During this task, the locations of the targets associated with small or large rewards and their corresponding delays were randomly varied. We found that prefrontal neurons often encoded the temporally discounted value of reward expected from a particular option. Furthermore, activity tended to increase with [corrected] discounted values for targets [corrected] presented in the neuron's preferred direction, suggesting that activity related to temporally discounted values in the prefrontal cortex might determine the animal's behavior during intertemporal choice.     

Ritual,emotion, and sacred symbols

This paper considers religion in relation to four recurrent traits: belief systems incorporating supernatural agents and counterintuitive concepts, communal ritual, separation of the sacred and the profane, and adolescence as a preferred developmental period for religious transmission. These co-occurring traits are viewed as an adaptive complex that offers clues to the evolution of religion from its nonhuman ritual roots. We consider the critical element differentiating religious from non-human ritual to be the conditioned association of emotion and abstract symbols. We propose neurophysiological mechanisms underlying such associations and argue that the brain plasticity of human adolescence constitutes an “experience expectant” developmental period for ritual conditioning of sacred symbols. We suggest that such symbols evolved to solve an ecological problem by extending communication and coordination of social relations across time and space. Candace Alcorta is currently a doctoral student in the Department of Anthropology at the University of Connecticut. Her research interests include the behavioral ecology and evolution of religion, and the interrelationship between cultural and neurophysiological systems. She is currently conducting research on adolescent religious participation, stress, and health. Richard Sosis is an associate professor in the Department of Anthropology at the University of Connecticut and a senior lecturer in the Department of Sociology and Anthropology at The Hebrew University of Jerusalem. His current research interests include the evolution of cooperation, utopian societies, and the behavioral ecology of religion. He has conducted fieldwork on Ifaluk Atoll in the Federated States of Micronesia and is currently pursuing various projects in Israel aimed at understanding the benefits and costs associated with religious behavior.     

Risk aversion and religious behaviour: Analysis using a sample of Danish twins

Economics offers an analytical framework to consider human behaviour including religious behaviour. Within the realm of Expected Utility Theory, religious belief and activity could be interpreted as an insurance both for current life events and for afterlife rewards. Based on that framework, we would expect that risk averse individuals would demand a more generous protection plan which they may do by devoting more effort and resources into religious activities such as church attendance and prayer, which seems to be in accordance with previous empirical results. However, a general concern regards the problems of spurious correlations due to underlying omitted or unobservable characteristics shaping both religious activities and risk attitudes. This paper examines empirically the demand for religion by analysing the association between risk attitudes on the one hand, and church attandance and prayer frequency on the other controlling for unobservable variables using survey data of Danish same-sex twin pairs. We verify the correlation between risk preferences and religion found previously by carrying out cross-sectional analyses. We also show that the association between risk attitudes and religious behaviour is driven by the subgroup of individuals who believe in an afterlife. In addition, when re-analysing our results using panel data analyses which cancel out shared factors among twin pairs, we find that the correlation found between risk aversion and religious behaviour is no longer significant indicating that other factors might explain differences in religious behaviour. Caution is needed in the interpretation of our results as the insignificant association between risk aversion and religious behaviour in the panel data analyses potentially might be due to measurement error causing attenuation bias or lack of variation within twin pairs rather than the actual absence of an association.     

Religion Does Matter for Climate Change Attitudes and Behavior

Little research has focused on the relationship between religion and climate change attitudes and behavior. Further, while there have been some studies examining the relationship between environmental attitudes and religion, most are focused on Christian denominations and secularism, and few have examined other religions such as Buddhism. Using an online survey of 1,927 Australians we examined links between membership of four religious groupings (Buddhists, Christian literalists and non-literalists, and Secularists) and climate change attitudes and behaviors. Differences were found across religious groups in terms of their belief in: (a) human induced climate change, (b) the level of consensus among scientists, (c) their own efficacy, and (d) the need for policy responses. We show, using ordinal regression, that religion explains these differences even after taking into account socio-demographic factors, knowledge and environmental attitude, including belief in man’s dominion over nature. Differences in attitude and behavior between these religious groups suggest the importance of engaging denominations to encourage change in attitudes and behavior among their members.     

Single units in the pigeon brain integrate reward amount and time-to-reward in an impulsive choice task

BACKGROUND: Animals prefer small over large rewards when the delays preceding large rewards exceed an individual tolerance limit. Such impulsive choice behavior occurs even in situations in which alternative strategies would yield more optimal outcomes. Behavioral research has shown that an animal's choice is guided by the alternative rewards' subjective values, which are a function of reward amount and time-to-reward. Despite increasing knowledge about the pharmacology and anatomy underlying impulsivity, it is still unknown how the brain combines reward amount and time-to-reward information to represent subjective reward value. RESULTS: We trained pigeons to choose between small, immediate rewards and large rewards delivered after gradually increasing delays. Single-cell recordings in the avian Nidopallium caudolaterale, the presumed functional analog of the mammalian prefrontal cortex, revealed that neural delay activation decreased with increasing delay length but also covaried with the expected reward amount. This integrated neural response was modulated by reward amount and delay, as predicted by a hyperbolical equation, of subjective reward value derived from behavioral studies. Furthermore, the neural activation pattern reflected the current reward preference and the time point of the shift from large to small rewards. CONCLUSIONS: The reported activity was modulated by the temporal devaluation of the anticipated reward in addition to reward amount. Our findings contribute to the understanding of neuropathologies such as drug addiction, pathological gambling, frontal lobe syndrome, and attention-deficit disorders, which are characterized by inappropriate temporal discounting and increased impulsiveness.     

Time Devours Things: How Impulsivity and Time Affect Temporal Decisions in Pathological Gamblers

Impulsivity is associated with several psychiatric disorders in which the loss of control of a specific behavior determines the syndrome itself. One particularly interesting population characterized by reported high impulsivity and problematic decision-making are those diagnosed with pathological gambling. However the association between impulsivity and decision making in pathological gambling has been only partially confirmed until now. We tested 23 normal controls and 23 diagnosed pathological gamblers in an intertemporal choice task, as well as other personality trait measurements. Results showed that gamblers scored higher on impulsivity questionnaires, and selected a higher percentage of impatient choices (higher percentage of smaller, sooner rewards), when compared to normal controls. Moreover, gamblers were faster in terms of reaction times at selecting the smaller, sooner options and discounted rewards more rapidly over time. Importantly, regression analyses clarified that self-reported measures of impulsivity played a significant role in biasing decisions towards small but more rapidly available rewards. In the present study we found evidence for impulsivity in personality traits and decisions in pathological gamblers relative to controls. We conclude by speculating on the need to incorporate impulsivity and decision biases in the conceptualization of pathological gambling for a better understanding and treatment of this pathology.     

Revisiting the Relationship between Individual Differences in Analytic Thinking and Religious Belief: Evidence That Measurement Order Moderates Their Inverse Correlation

Prior research has found that persons who favor more analytic modes of thought are less religious. We propose that individual differences in analytic thought are associated with reduced religious beliefs particularly when analytic thought is measured (hence, primed) first. The current study provides a direct replication of prior evidence that individual differences in analytic thinking are negatively related to religious beliefs when analytic thought is measured before religious beliefs. When religious belief is measured before analytic thinking, however, the negative relationship is reduced to non-significance, suggesting that the link between analytic thought and religious belief is more tenuous than previously reported. The current study suggests that whereas inducing analytic processing may reduce religious belief, more analytic thinkers are not necessarily less religious. The potential for measurement order to inflate the inverse correlation between analytic thinking and religious beliefs deserves additional consideration.     

Why Do You Believe in God? Relationships between Religious Belief,Analytic Thinking,Mentalizing and Moral Concern

Prior work has established that analytic thinking is associated with disbelief in God, whereas religious and spiritual beliefs have been positively linked to social and emotional cognition. However, social and emotional cognition can be subdivided into a number of distinct dimensions, and some work suggests that analytic thinking is in tension with some aspects of social-emotional cognition. This leaves open two questions. First, is belief linked to social and emotional cognition in general, or a specific dimension in particular? Second, does the negative relationship between belief and analytic thinking still hold after relationships with social and emotional cognition are taken into account? We report eight hypothesis-driven studies which examine these questions. These studies are guided by a theoretical model which focuses on the distinct social and emotional processing deficits associated with autism spectrum disorders (mentalizing) and psychopathy (moral concern). To our knowledge no other study has investigated both of these dimensions of social and emotion cognition alongside analytic thinking. We find that religious belief is robustly positively associated with moral concern (4 measures), and that at least part of the negative association between belief and analytic thinking (2 measures) can be explained by a negative correlation between moral concern and analytic thinking. Using nine different measures of mentalizing, we found no evidence of a relationship between mentalizing and religious or spiritual belief. These findings challenge the theoretical view that religious and spiritual beliefs are linked to the perception of agency, and suggest that gender differences in religious belief can be explained by differences in moral concern. These findings are consistent with the opposing domains hypothesis, according to which brain areas associated with moral concern and analytic thinking are in tension.     

The evolution of social discounting in hierarchically clustered populations

The expression of a social behaviour may affect the fitness of actors and recipients living in the present and in the future of the population. When there is a risk that a future reward will not be experienced in such a context, the value of that reward should be discounted; but by how much? Here, we evaluate social discount rates for delayed fitness rewards to group of recipients living at different positions in both space and time than the actor in a hierarchically clustered population. This is a population where individuals are grouped into families, families into villages, villages into clans, and so on, possibly ad infinitum. The group-wide fitness effects are assumed to either increase or decrease the fecundity or the survival of recipients and can be arbitrarily extended in space and time. We find that actions changing the survival of individuals living in the future are generally more strongly discounted than fecundity-changing actions for all future times and that the value of future rewards increases as individuals live longer. We also find that delayed fitness effects may not only be discounted by a constant factor per unit delay (exponential discounting), but that, as soon as there is localized dispersal in a population, discounting per unit delay is likely to fall rapidly for small delays and then slowly for longer delays (hyperbolic discounting). As dispersal tends to be localized in natural populations, our results suggest that evolution is likely to favour individuals that express present-biased behaviours and that may be time-inconsistent with respect to their group-wide effects.     

Within-subject differences in degree of delay discounting as a function of order of presentation of hypothetical cash rewards

Procedural variants in estimating delay discounting (DD) have been shown to yield significant differences in estimated degree of DD as well as variations in individual patterns of choice. For example, a recent study found significantly different degrees of DD between groups assessed using either an ascending or descending order of presentation of the immediately available rewards. The purpose of this study was to test for within-subject effects of order of presentation of the immediate rewards in a DD task. In a single session, college students (N = 29) were asked to complete two DD tasks, one with the immediate rewards presented in ascending order and one in descending order. Consistent with previous results, significantly larger mean area under the discounting curve (AUC) was observed when the descending sequence was used compared to the ascending order of presentation; and the correlation between both measurements was moderate. These results suggest that some DD assessment tasks may be sensitive to contextual variables such as order and range of the reward and delay values.     

Exponential Versus Hyperbolic Discounting of Delayed Outcomes: Risk and Waiting Time

Frequently, animals must choose between more immediate, smallerrewards and more delayed, but larger rewards. For example, theyoften must decide between accepting a smaller prey item versuscontinuing to search for a larger one, or between entering aleaner patch versus travelling to a richer patch that is furtheraway. In both situations, choice of the more immediate, butsmaller reward may be interpreted as implying that the valueof the later reward is discounted; that is, the value of thelater reward decreases as the delay to its receipt increases.This decrease in value may occur because of the increased riskinvolved in waiting for rewards, or because of the decreasedrate of reward associated with increased waiting time. The presentresearch attempts to determine the form of the relation betweenvalue and delay, and examines implications of this relationfor mechanisms underlying risk-sensitive foraging. Two accounts of the relation between value and delay have beenproposed to describe the decrease in value resulting from increasesin delay: an exponential model and a hyperbolic model. Our researchdemonstrates that, of the two, a hyperbola-like discountingmodel consistently explains more of the variance in temporaldiscounting data at the group level and, importantly, at theindividual level as well. We show mathematically that the hyperbolicmodel shares fundamental features with models of prey and patchchoice. In addition, the present review highlights the implicationsof a psychological perspective for the behavioral biology ofrisksensitive foraging, as well as the implications of an ecologicalperspective for the behavioral psychology of risk-sensitivechoice and decision-making.     

Adolescent religiousness and its influence on substance use: preliminary findings from the Mid-Atlantic School Age Twin Study.

Research has consistently shown that religiousness is associated with lower levels of alcohol and drug use, but little is known about the nature of adolescent religiousness or the mechanisms through which it influences problem behavior in this age group. This paper presents preliminary results from the Mid-Atlantic School Age Twin Study, a prospective, population-based study of 6-18-year-old twins and their mothers. Factor analysis of a scale developed to characterize adolescent religiousness, the Religious Attitudes and Practices Inventory (RAPI), revealed three factors: theism, religious/spiritual practices, and peer religiousness. Twin correlations and univariate behavior-genetic models for these factors and a measure of belief that drug use is sinful reveal in 357 twin pairs that common environmental factors significantly influence these traits, but a minor influence of genetic factors could not be discounted. Correlations between the multiple factors of adolescent religiousness and substance use, comorbid problem behavior, mood disorders, and selected risk factors for substance involvement are also presented. Structural equation modeling illustrates that specific religious beliefs about the sinfulness of drugs and level of peer religiousness mediate the relationship between theistic beliefs and religious/spiritual practices on substance use. Limitations and future analyses are discussed.     

Corvids can decide if a future exchange is worth waiting for

Evidence for time-dependent calculations about future rewards is scarce in non-human animals. In non-human primates, only great apes are comparable with humans. Still, some species wait for several minutes to obtain a better reward in delayed exchange tasks. Corvids have been shown to match with non-human primates in some time-related tasks. Here, we investigate a delay of gratification in two corvid species, the carrion crow (Corvus corone) and the common raven (Corvus corax), in an exchange task. Results show that corvids success decreases quickly as delay increases, with a maximal delay of up to 320 s (more than 5 min). The decision to wait rests both on the quality of the prospective reward and the time required to obtain it. Corvids also apply tactics (placing the reward on the ground or caching it) that probably alleviate costs of waiting and distract their attention during waiting. These findings contrast previous results on delayed gratification in birds and indicate that some species may perform comparably to primates.