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Learning to respond optimally under a broad array of environmental conditions is a critical brain function that requires engaging the cognitive systems that are optimal for solving the task at hand. Serotonin is implicated in learning and decision-making, but the specific functions of serotonin in system-level cognitive control remain unclear. Across 3 studies, we examined the influence of a polymorphism within the promoter region of the serotonin transporter gene (5-HTTLPR polymorphism in SLC6A4) on participants' ability to engage the task appropriate cognitive system when the reflexive (Experiments 1 and 2) or the reflective (Experiment 3) system was optimal. Critically, we utilized a learning task for which all aspects remain fixed with only the nature of the optimal cognitive processing system varying across experiments. Using large community samples, Experiments 1 and 2 (screened for psychiatric diagnosis) found that 5-HTTLPR S/LG allele homozygotes, with putatively lower serotonin transport functionality, outperformed LA allele homozygotes in a reflexive-optimal learning task. Experiment 3 used a large community sample, also screened for psychiatric diagnosis, and found that 5-HTTLPR LA homozygotes, with putatively higher serotonin transport functionality, outperformed S/LG allele homozygotes in a reflective-optimal learning task.

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Although research on aging and decision making continues to grow, the majority of studies examine decisions made to maximize monetary earnings or points. It is not clear whether these results generalize to other types of rewards. To investigate this, we examined adult age differences in 92 healthy participants aged 22 to 83. Participants completed 9 hypothetical discounting tasks, which included 3 types of discounting factors (time, probability, effort) across 3 reward domains (monetary, social, health). Participants made choices between a smaller magnitude reward with a shorter time delay/higher probability/lower level of physical effort required and a larger magnitude reward with a longer time delay/lower probability/higher level of physical effort required. Older compared with younger individuals were more likely to choose options that involved shorter time delays or higher probabilities of experiencing an interaction with a close social partner or receiving health benefits from a hypothetical drug. These findings suggest that older adults may be more motivated than young adults to obtain social and health rewards immediately and with certainty. (PsycINFO Database Record

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Older and younger adults performed a state-based decision-making task while undergoing functional MRI (fMRI). We proposed that younger adults would be more prone to base their decisions on expected value comparisons, but that older adults would be more reactive decision-makers who would act in response to recent changes in rewards or states, rather than on a comparison of expected values. To test this we regressed BOLD activation on two measures from a sophisticated reinforcement learning (RL) model. A value-based regressor was computed by subtracting the immediate value of the selected alternative from its long-term value. The other regressor was a state-change uncertainty signal that served as a proxy for whether the participant's state improved or declined, relative to the previous trial. Younger adults' activation was modulated by the value-based regressor in ventral striatal and medial PFC regions implicated in reinforcement learning. Older adults' activation was modulated by state-change uncertainty signals in right dorsolateral PFC, and activation in this region was associated with improved performance in the task. This suggests that older adults may depart from standard expected-value based strategies and recruit lateral PFC regions to engage in reactive decision-making strategies.

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Age-related deficits are seen across tasks where learning depends on asocial feedback processing, however plasticity has been observed in some of the same tasks in social contexts suggesting a novel way to attenuate deficits. Socioemotional selectivity theory suggests this plasticity is due to a deliberative motivational shift toward achieving well-being with age (positivity effect) that reverses when executive processes are limited (negativity effect). The present study examined the interaction of feedback valence (positive, negative) and social salience (emotional face feedback - happy; angry, asocial point feedback - gain; loss) on learning in a deliberative task that challenges executive processes and a procedural task that does not. We predict that angry face feedback will improve learning in a deliberative task when executive function is challenged. We tested two competing hypotheses regarding the interactive effects of deliberative emotional biases on automatic feedback processing: (1) If deliberative emotion regulation and automatic feedback are interactive we expect happy face feedback to improve learning and angry face feedback to impair learning in older adults because cognitive control is available. (2) If deliberative emotion regulation and automatic feedback are not interactive we predict that emotional face feedback will not improve procedural learning regardless of valence. Results demonstrate that older adults show persistent deficits relative to younger adults during procedural category learning suggesting that deliberative emotional biases do not interact with automatic feedback processing. Interestingly, a subgroup of older adults identified as potentially using deliberative strategies tended to learn as well as younger adults with angry relative to happy feedback, matching the pattern observed in the deliberative task. Results suggest that deliberative emotional biases can improve deliberative learning, but have no effect on procedural learning.

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Humans with seven or more repeats in exon III of the DRD4 gene (long DRD4 carriers) sometimes demonstrate impaired attention, as seen in attention-deficit hyperactivity disorder, and at other times demonstrate heightened attention, as seen in addictive behavior. Although the clinical effects of DRD4 are the focus of much work, this gene may not necessarily serve as a "risk" gene for attentional deficits, but as a plasticity gene where attention is heightened for priority items in the environment and impaired for minor items. Here we examine the role of DRD4 in two tasks that benefit from selective attention to high-priority information. We examine a category learning task where performance is supported by focusing on features and updating verbal rules. Here, selective attention to the most salient features is associated with good performance. In addition, we examine the Operation Span (OSPAN) task, a working memory capacity task that relies on selective attention to update and maintain items in memory while also performing a secondary task. Long DRD4 carriers show superior performance relative to short DRD4 homozygotes (six or less tandem repeats) in both the category learning and OSPAN tasks. These results suggest that DRD4 may serve as a "plasticity" gene where individuals with the long allele show heightened selective attention to high-priority items in the environment, which can be beneficial in the appropriate context.

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Recent decision-making work has focused on a distinction between a habitual, model-free neural system that is motivated toward actions that lead directly to reward and a more computationally demanding goal-directed, model-based system that is motivated toward actions that improve one's future state. In this article, we examine how aging affects motivation toward reward-based versus state-based decision making. Participants performed tasks in which one type of option provided larger immediate rewards but the alternative type of option led to larger rewards on future trials, or improvements in state. We predicted that older adults would show a reduced preference for choices that led to improvements in state and a greater preference for choices that maximized immediate reward. We also predicted that fits from a hybrid reinforcement-learning model would indicate greater model-based strategy use in younger than in older adults. In line with these predictions, older adults selected the options that maximized reward more often than did younger adults in three of the four tasks, and modeling results suggested reduced model-based strategy use. In the task where older adults showed similar behavior to younger adults, our model-fitting results suggested that this was due to the utilization of a win-stay-lose-shift heuristic rather than a more complex model-based strategy. Additionally, within older adults, we found that model-based strategy use was positively correlated with memory measures from our neuropsychological test battery. We suggest that this shift from state-based to reward-based motivation may be due to age related declines in the neural structures needed for more computationally demanding model-based decision making.

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Depression is often characterized by attentional biases toward negative items and away from positive items, which likely affects reward and punishment processing. Recent work has reported that training attention away from negative stimuli reduced this bias and reduced depressive symptoms. However, the effect of attention training on subsequent learning has yet to be explored. In the present study, participants were required to learn to maximize reward during decision making. Undergraduates with elevated self-reported depressive symptoms received attention training toward positive stimuli prior to performing the decision-making task (n = 20; active training). The active-training group was compared to two other groups: undergraduates with elevated self-reported depressive symptoms who received placebo training (n = 22; placebo training) and a control group with low levels of depressive symptoms (n = 33; nondepressive control). The placebo-training depressive group performed worse and switched between options more than did the nondepressive controls on the reward maximization task. However, depressives that received active training performed as well as the nondepressive controls. Computational modeling indicated that the placebo-trained group learned more from negative than from positive prediction errors, leading to more frequent switching. The nondepressive control and active-training depressive groups showed similar learning from positive and negative prediction errors, leading to less-frequent switching and better performance. Our results indicate that individuals with elevated depressive symptoms are impaired at reward maximization, but that the deficit can be improved with attention training toward positive stimuli.

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We examined the relationship between pressure and age-related changes in decision-making using a task for which currently available rewards depend on the participant's previous history of choices. Optimal responding in this task requires the participant to learn how his or her current choices affect changes in the future rewards given for each option. Building on the scaffolding theory of aging and cognition, we predicted that when additional frontal resources are available, compensatory recruitment leads to increased monitoring and increased use of heuristic-based strategies, ultimately leading to better performance. Specifically, we predicted that scaffolding would result in an age-related performance advantage under no pressure conditions. We also predicted that, although younger adults would engage in scaffolding under pressure, older adults would not have additional resources available for increased scaffolding under pressure-packed conditions, leading to an age-related performance deficit. Both predictions were supported by the data. In addition, computational models were used to evaluate decision-making strategies employed by each participant group. As expected, older adults under no pressure conditions and younger adults under pressure conditions showed increased use of heuristic-based strategies relative to older adults under pressure and younger adults under no pressure, respectively. These results are consistent with the notion that scaffolding can occur across the life span in the face of an environmental challenge.

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Arousal Biased Competition theory suggests that arousal enhances competitive attentional processes, but makes no strong claims about valence effects. Research suggests that the scope of enhanced attention depends on valence with negative arousal narrowing and positive arousal broadening attention. Attentional scope likely affects declarative-memory-mediated and perceptual-representation-mediated learning systems differently, with declarative-memory-mediated learning depending on narrow attention to develop targeted verbalizable rules, and perceptual-representation-mediated learning depending on broad attention to develop a perceptual representation. We hypothesize that negative arousal accentuates declarative-memory-mediated learning and attenuates perceptual-representation-mediated learning, while positive arousal reverses this pattern. Prototype learning provides an ideal test bed as dissociable declarative-memory and perceptual-representation systems mediate two-prototype (AB) and one-prototype (AN) prototype learning, respectively, and computational models are available that provide powerful insights on cognitive processing. As predicted, we found that negative arousal narrows attentional focus facilitating AB learning and impairing AN learning, while positive arousal broadens attentional focus facilitating AN learning and impairing AB learning.

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Animal research and human neuroimaging studies indicate that stress increases dopamine levels in brain regions involved in reward processing, and stress also appears to increase the attractiveness of addictive drugs. The current study tested the hypothesis that stress increases reward salience, leading to more effective learning about positive than negative outcomes in a probabilistic selection task. Changes to dopamine pathways with age raise the question of whether stress effects on incentive-based learning differ by age. Thus, the present study also examined whether effects of stress on reinforcement learning differed for younger (age 18-34) and older participants (age 65-85). Cold pressor stress was administered to half of the participants in each age group, and salivary cortisol levels were used to confirm biophysiological response to cold stress. After the manipulation, participants completed a probabilistic learning task involving positive and negative feedback. In both younger and older adults, stress enhanced learning about cues that predicted positive outcomes. In addition, during the initial learning phase, stress diminished sensitivity to recent feedback across age groups. These results indicate that stress affects reinforcement learning in both younger and older adults and suggests that stress exerts different effects on specific components of reinforcement learning depending on their neural underpinnings.

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Previous research reveals that older adults sometimes show enhanced processing of emotionally positive stimuli relative to negative stimuli, but that this positivity bias reverses to become a negativity bias when cognitive control resources are less available. In this study, we test the hypothesis that emotionally positive feedback will attenuate well-established age-related deficits in rule learning whereas emotionally negative feedback will amplify age deficits-but that this pattern will reverse when the task involves a high cognitive load. Experiment 1 used emotional face feedback and revealed an interaction among age, valence of the feedback, and task load. When the task placed minimal load on cognitive control resources, happy-face feedback attenuated age-related deficits in initial rule learning and angry-face feedback led to age-related deficits in initial rule learning and set shifting. However, when the task placed a high load on cognitive control resources, we found that angry-face feedback attenuated age-related deficits in initial rule learning and set shifting whereas happy-face feedback led to age-related deficits in initial rule learning and set shifting. Experiment 2 used less emotional point feedback and revealed age-related deficits in initial rule learning and set shifting under low and high cognitive load for point-gain and point-loss conditions. The research presented here demonstrates that emotional feedback can attenuate age-related learning deficits-but only positive feedback for tasks with a low cognitive load and negative feedback for tasks with high cognitive load.

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Prior research indicates that depressed individuals are less responsive to rewards and more sensitive to punishments than non-depressed individuals. This study examines decision-making under reward maximizing or punishment minimizing conditions among adults with low (n=47) or high (n=48) depression symptoms. We utilized a history-independent decision-making task where learning is experience-based and the participants' goal is to enhance immediate payoff. Results indicated a significant interaction between incentive condition (reward maximizing, punishment minimizing) and depression group. Within the low depression group, better performance was observed for reward maximization than punishment minimization. In contrast, within the high depression group, better performance was observed for punishment minimization than reward maximization. Further, the high depression group outperformed the low depression symptom group in the punishment minimization condition, but no depression group differences were observed in the reward maximization condition. Computational modeling indicated that the high depression group was more likely to choose options with the highest expected reward, particularly in the punishment condition. Thus, decision-making is improved for people with elevated depression symptoms when minimizing punishment relative to maximizing rewards.

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A prevalent stereotype is that people become less risk taking and more cautious as they get older. However, in laboratory studies, findings are mixed and often reveal no age differences. In the current series of experiments, we examined whether age differences in risk seeking are more likely to emerge when choices include a certain option (a sure gain or a sure loss). In four experiments, we found that age differences in risk preferences only emerged when participants were offered a choice between a risky and a certain gamble but not when offered two risky gambles. In particular, Experiments 1 and 2 included only gambles about potential gains. Here, compared with younger adults, older adults preferred a certain gain over a chance to win a larger gain and thus, exhibited more risk aversion in the domain of gains. But in Experiments 3 and 4, when offered the chance to take a small sure loss rather than risking a larger loss, older adults exhibited more risk seeking in the domain of losses than younger adults. Both their greater preference for sure gains and greater avoidance of sure losses suggest that older adults weigh certainty more heavily than younger adults. Experiment 4 also indicates that older adults focus more on positive emotions than younger adults do when considering their options, and that this emotional shift can at least partially account for age differences in how much people are swayed by certainty in their choices.

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Individuals with depressive symptoms typically show deficits in decision-making. However, most work has emphasized decision-making under gain-maximization conditions. A gain-maximization framework may undermine decision-making when depressive symptoms are present because depressives are generally more sensitive to losses than gains. The present study examined decision making in a non-clinical sample of depressive and non-depressive individuals under gain-maximization or loss-minimization conditions using a task for which the currently available rewards depend upon participants' previous history of choices. As predicted, we found a cross over interaction whereby depressive individuals were superior to non-depressive individuals under loss-minimization conditions, but were inferior to non-depressive individuals under gain-maximization conditions. In addition, we found that loss-minimization performance was superior to gain-maximization performance for depressive individuals, but that gain-maximization performance was superior to loss-minimization performance for non-depressive individuals. These results suggest that decision making deficits observed when depressive symptoms are present may be attenuated when decisions involve minimizing losses rather than maximizing gains.

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Past studies have revealed that encountering negative events interferes with cognitive processing of subsequent stimuli. The present study investigates whether negative events affect semantic and perceptual processing differently. Presentation of negative pictures produced slower reaction times than neutral or positive pictures in tasks that require semantic processing, such as natural or man-made judgments about drawings of objects, commonness judgments about objects, and categorical judgments about pairs of words. In contrast, negative picture presentation did not slow down judgments in subsequent perceptual processing (e.g., color judgments about words, size judgments about objects). The subjective arousal level of negative pictures did not modulate the interference effects on semantic or perceptual processing. These findings indicate that encountering negative emotional events interferes with semantic processing of subsequent stimuli more strongly than perceptual processing, and that not all types of subsequent cognitive processing are impaired by negative events.

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In two experiments, younger and older adults performed decision-making tasks in which reward values available were either independent of or dependent on the previous sequence of choices made. The choice-independent task involved learning and exploiting the options that gave the highest rewards on each trial. In this task, the stability of the expected reward for each option was not influenced by the previous choices participants made. The choice-dependent task involved learning how each choice influenced future rewards for two options and making the best decisions based on that knowledge. Younger adults performed better when rewards were independent of choice, whereas older adults performed better when rewards were dependent on choice. These findings suggest a fundamental difference in the way in which younger adults and older adults approach decision-making situations. We discuss the results in the context of prominent decision-making theories and offer possible explanations based on neurobiological and behavioral changes associated with aging.

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In a typical reversal-learning experiment, one learns stimulus-outcome contingencies that then switch without warning. For instance, participants might have to repeatedly choose between two faces, one of which yields points whereas the other does not, with a reversal at some point in which face yields points. The current study examined age differences in the effects of outcome type on reversal learning. In the first experiment, the participants' task was either to select the person who would be in a better mood or to select the person who would yield more points. Reversals in which face was the correct option occurred several times. Older adults did worse in blocks in which the correct response was to select the person who would not be angry than in blocks in which the correct response was to select the person who would smile. Younger adults did not show a difference by emotional valence. In the second study, the negative condition was switched to have the same format as the positive condition (to select who will be angry). Again, older adults did worse with negative than positive outcomes, whereas younger adults did not show a difference by emotional valence. A third experiment replicated the lack of valence effects in younger adults with a harder probabilistic reversal-learning task. In the first two experiments, older adults performed about as well as younger adults in the positive conditions but performed worse in the negative conditions. These findings suggest that negative emotional outcomes selectively impair older adults' reversal learning.

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Under stress, men tend to withdraw socially whereas women seek social support. This functional magnetic resonance imaging study indicates that stress also affects brain activity while viewing emotional faces differently for men and women. Fusiform face area response to faces was diminished by acute stress in men but increased by stress in women. Furthermore, among stressed men viewing angry faces, brain regions involved in interpreting and understanding others' emotions (the insula, temporal pole, and inferior frontal gyrus) showed reduced coordination with the fusiform face area and the amygdala, whereas the functional connectivity among these regions increased with stress for women. These findings suggest that stress influences emotional perception differently for men and women.

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Showing an arousing central stimulus in a scene often leads to enhanced memory for the arousing central information and impaired memory for peripheral details. However, it is not clear from previous work whether arousing stimuli impair memory for all nonarousing nearby information or just background information. In several experiments, we tested how emotionally arousing pictures affect memory for nearby pictures and for background information. We found that when 2 pictures were presented together, an arousing picture did not affect item and location memory for the other picture. In contrast, an arousing picture impaired memory for a background pattern. These findings suggest that arousal impairs memory for information that is the target of perceptual suppression, such as background information when there is a figure-ground distinction, but does not impair memory for other foreground information.

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BACKGROUND: Decisions involving risk often must be made under stressful circumstances. Research on behavioral and brain differences in stress responses suggest that stress might have different effects on risk taking in males and females. METHODOLOGY/PRINCIPAL FINDINGS: In this study, participants played a computer game designed to measure risk taking (the Balloon Analogue Risk Task) fifteen minutes after completing a stress challenge or control task. Stress increased risk taking among men but decreased it among women. CONCLUSIONS/SIGNIFICANCE: Acute stress amplifies sex differences in risk seeking; making women more risk avoidant and men more risk seeking. Evolutionary principles may explain these stress-induced sex differences in risk taking behavior.

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