A polygenic score for educational attainment partially predicts voter turnout.

Twin and adoption studies have shown that individual differences in political participation can be explained, in part, by genetic variation. However, these research designs cannot identify which genes are related to voting or the pathways through which they exert influence, and their conclusions rely on possibly restrictive assumptions. In this study, we use three different US samples and a Swedish sample to test whether genes that have been identified as associated with educational attainment, one of the strongest correlates of political participation, predict self-reported and validated voter turnout. We find that a polygenic score capturing individuals' genetic propensity to acquire education is significantly related to turnout. The strongest associations we observe are in second-order midterm elections in the United States and European Parliament elections in Sweden, which tend to be viewed as less important by voters, parties, and the media and thus present a more information-poor electoral environment for citizens to navigate. A within-family analysis suggests that individuals' education-linked genes directly affect their voting behavior, but, for second-order elections, it also reveals evidence of genetic nurture. Finally, a mediation analysis suggests that educational attainment and cognitive ability combine to account for between 41% and 63% of the relationship between the genetic propensity to acquire education and voter turnout.

Conscientiousness associated with efficiency of the salience/ventral attention network: Replication in three samples using individualized parcellation.

Conscientiousness, and related constructs impulsivity and self-control, have been related to structural and functional properties of regions in the prefrontal cortex (PFC) and anterior insula. Network-based conceptions of brain function suggest that these regions belong to a single large-scale network, labeled the salience/ventral attention network (SVAN). The current study tested associations between conscientiousness and resting-state functional connectivity in this network using two community samples (N's = 244 and 239) and data from the Human Connectome Project (N = 1000). Individualized parcellation was used to improve functional localization accuracy and facilitate replication. Functional connectivity was measured using an index of network efficiency, a graph theoretical measure quantifying the capacity for parallel information transfer within a network. Efficiency of a set of parcels in the SVAN was significantly associated with conscientiousness in all samples. Findings are consistent with a theory of conscientiousness as a function of variation in neural networks underlying effective prioritization of goals.

Risky decisions are influenced by individual attributes as a function of risk preference.

It has long been assumed in economic theory that multi-attribute decisions involving several attributes or dimensions - such as probabilities and amounts of money to be earned during risky choices - are resolved by first combining the attributes of each option to form an overall expected value and then comparing the expected values of the alternative options, using a unique evidence accumulation process. A plausible alternative would be performing independent comparisons between the individual attributes and then integrating the results of the comparisons afterwards. Here, we devise a novel method to disambiguate between these types of models, by orthogonally manipulating the expected value of choice options and the relative salience of their attributes. Our results, based on behavioral measures and drift-diffusion models, provide evidence in favor of the framework where information about individual attributes independently impacts deliberation. This suggests that risky decisions are resolved by running in parallel multiple comparisons between the separate attributes - possibly alongside an additional comparison of expected value. This result stands in contrast with the assumption of standard economic theory that choices require a unique comparison of expected values and suggests that at the cognitive level, decision processes might be more distributed than commonly assumed. Beyond our planned analyses, we also discovered that attribute salience affects people of different risk preference type in different ways: risk-averse participants seem to focus more on probability, except when monetary amount is particularly high; risk-neutral/seeking participants, in contrast, seem to focus more on monetary amount, except when probability is particularly low.

Theory matters for identifying a causal role for genetic factors in socioeconomic outcomes.

Any empirical claim about the role of genes in socioeconomic outcomes involves successfully addressing the identification problem. This commentary argues that socioeconomic outcomes such as education are sufficiently complex, involving so many mechanisms, that understanding the role genes requires the use of formal theoretical structures.

The association between saving disposition and financial distress: A genetically informed approach.

Saving disposition, the tendency to save rather than consume, has been found to be associated with economic outcomes. People lacking the disposition to save are more likely to experience financial distress. This association could be driven by other economic factors, behavioral traits, or even genetic effects. Using a sample of 3,920 American twins, we develop scales to measure saving disposition and financial distress. We find genetic influences on both traits, but also a large effect of the rearing family environment on saving disposition. We estimate that 44% of the covariance between the two traits is due to genetic effects. Saving disposition remains strongly associated with lower financial distress, even after controlling for family income, cognitive ability, and personality traits. The association persists within families and monozygotic twin pairs; the twin who saves more tends to be the twin who experiences less financial distress. This result suggest that there is a direct association between saving disposition and financial distress, although the direction of causation remains unclear.

The role of parental genotype in predicting offspring years of education: evidence for genetic nurture.

Similarities between parent and offspring are widespread in psychology; however, shared genetic variants often confound causal inference for offspring outcomes. A polygenic score (PGS) derived from genome-wide association studies (GWAS) can be used to test for the presence of parental influence that controls for genetic variants shared across generations. We use a PGS for educational attainment (EA3; N ≈ 750 thousand) to predict offspring years of education in a sample of 2517 twins and both parents. We find that within families, the dizygotic twin with the higher PGS is more likely to attain higher education (unstandardized ß = 0.32; p < 0.001). Additionally, however, we find an effect of parental genotype on offspring outcome that is independent of the offspring's own genotype; this raises the variance explained in offspring years of education from 9.3 to 11.1% (∆R2 = 0.018, p < 0.001). Controlling for parental IQ or socioeconomic status substantially attenuated or eliminated this effect of parental genotype. These findings suggest a role of environmental factors affected by heritable characteristics of the parents in fostering offspring years of education.

The Contribution of Cognitive and Noncognitive Skills to Intergenerational Social Mobility.

We investigated intergenerational educational and occupational mobility in a sample of 2,594 adult offspring and 2,530 of their parents. Participants completed assessments of general cognitive ability and five noncognitive factors related to social achievement; 88% were also genotyped, allowing computation of educational-attainment polygenic scores. Most offspring were socially mobile. Offspring who scored at least 1 standard deviation higher than their parents on both cognitive and noncognitive measures rarely moved down and frequently moved up. Polygenic scores were also associated with social mobility. Inheritance of a favorable subset of parent alleles was associated with moving up, and inheritance of an unfavorable subset was associated with moving down. Parents' education did not moderate the association of offspring's skill with mobility, suggesting that low-skilled offspring from advantaged homes were not protected from downward mobility. These data suggest that cognitive and noncognitive skills as well as genetic factors contribute to the reordering of social standing that takes place across generations.

Dissociation between Private and Social Counterfactual Value Signals Following Ventromedial Prefrontal Cortex Damage.

Individuals learn by comparing the outcome of chosen and unchosen actions. A negative counterfactual value signal is generated when this comparison is unfavorable. This can happen in private as well as in social settings-where the foregone outcome results from the choice of another person. We hypothesized that, despite sharing similar features such as supporting learning, these two counterfactual signals might implicate distinct brain networks. We conducted a neuropsychological study on the role of private and social counterfactual value signals in risky decision-making. Patients with lesions in the ventromedial prefrontal cortex (vmPFC), lesion controls, and healthy controls repeatedly chose between lotteries. In private trials, participants could observe the outcomes of their choices and the outcomes of the unselected lotteries. In social trials, participants could also see the other player's choices and outcome. At the time of outcome, vmPFC patients were insensitive to private counterfactual value signals, whereas their responses to social comparison were similar to those of control participants. At the time of choice, intact vmPFC was necessary to integrate counterfactual signals in decisions, although amelioration was observed during the course of the task, possibly driven by social trials. We conclude that if the vmPFC is critical in processing private counterfactual signals and in integrating those signals in decision-making, then distinct brain areas might support the processing of social counterfactual signals.

The goal priority network as a neural substrate of Conscientiousness.

Conscientiousness is a personality trait associated with many important life outcomes, but little is known about the mechanisms that underlie it. We investigated its neural correlates using functional connectivity analysis in fMRI, which identifies brain regions that act in synchrony. We tested the hypothesis that a broad network resembling a combination of the salience and ventral attention networks, which we provisionally label the goal priority network (GPN), is a neural correlate of Conscientiousness. Self- and peer-ratings of Conscientiousness were collected in a community sample of adults who underwent a resting-state fMRI scan (N = 218). An independent components analysis yielded five components that overlapped substantially with the GPN. We examined synchrony within and between these GPN subcomponents. Synchrony within one of the components-mainly comprising regions of anterior insula, dorsal anterior cingulate cortex, and dorsolateral prefrontal cortex-was significantly associated with Conscientiousness. Connectivity between this component and the four other GPN components was also significantly associated with Conscientiousness. Our results support the hypothesis that variation in a network that enables prioritization of multiple goals may be central to Conscientiousness.

Testosterone administration does not affect men's rejections of low ultimatum game offers or aggressive mood.

Correlative evidence suggests that testosterone promotes dominance and aggression. However, causal evidence is scarce and offers mixed results. To investigate this relationship, we administered testosterone for 48h to 41 healthy young adult men in a within-subjects, double-blind placebo-controlled balanced crossover design. Subjects played the role of responders in an ultimatum game, where rejecting a low offer is costly, but serves to destroy the proposer's profit. Such action can hence be interpreted as non-physical aggression in response to social provocation. In addition, subjects completed a self-assessed mood questionnaire. As expected, self-reported aggressiveness was a key predictor of ultimatum game rejections. However, while testosterone affected subjective ratings of feeling energetic and interested, our evidence strongly suggests that testosterone had no effect on ultimatum game rejections or on aggressive mood. Our findings illustrate the importance of using causal interventions to assess correlative evidence.

Cognitive, Noncognitive, and Family Background Contributions to College Attainment: A Behavioral Genetic Perspective.

There is considerable evidence that college attainment is associated with family background and cognitive and noncognitive skills. Behavioral genetic methods are used to determine whether the family background effect is mediated through cognitive and noncognitive skill development. We analyze data from two longitudinal behavioral genetic studies: the Minnesota Twin Family Study, consisting of 1,382 pairs of like-sex twins and their parents, and the Sibling Interaction and Behavior Study, consisting of 409 adoptive and 208 nonadoptive families with two offspring and their rearing parents. Cognitive ability, noncognitive skills, and family background are all associated with offspring college attainment. Biometric analysis shows that the intergenerational transmission of college attainment owes to both genetic and shared environmental factors. The shared environmental influence was not due to highly educated parents fostering noncognitive skill development in their children, and there was limited evidence that they foster cognitive skill development. The environmental transmission of educational attainment does not appear to be a consequence of highly educated parents fostering cognitive and noncognitive skill development. Alternative mechanisms are needed to explain the strong shared environmental influence on college attainment. Possibilities include academic expectations, social network effects, and the economic benefits of having wealthy parents.

Children's strategic theory of mind.

Human strategic interaction requires reasoning about other people's behavior and mental states, combined with an understanding of their incentives. However, the ontogenic development of strategic reasoning is not well understood: At what age do we show a capacity for sophisticated play in social interactions? Several lines of inquiry suggest an important role for recursive thinking (RT) and theory of mind (ToM), but these capacities leave out the strategic element. We posit a strategic theory of mind (SToM) integrating ToM and RT with reasoning about incentives of all players. We investigated SToM in 3- to 9-y-old children and adults in two games that represent prevalent aspects of social interaction. Children anticipate deceptive and competitive moves from the other player and play both games in a strategically sophisticated manner by 7 y of age. One game has a pure strategy Nash equilibrium: In this game, children achieve equilibrium play by the age of 7 y on the first move. In the other game, with a single mixed-strategy equilibrium, children's behavior moved toward the equilibrium with experience. These two results also correspond to two ways in which children's behavior resembles adult behavior in the same games. In both games, children's behavior becomes more strategically sophisticated with age on the first move. Beyond the age of 7 y, children begin to think about strategic interaction not myopically, but in a farsighted way, possibly with a view to cooperating and capitalizing on mutual gains in long-run relationships.

When gaze opens the channel for communication: Integrative role of IFG and MPFC.

Recent advances in the field of cognitive neuroscience have revealed that direct gaze modulates activity in cortical and subcortical key regions of the 'social brain network', including the inferior frontal gyrus (IFG) and the anterior rostral medial prefrontal cortex (arMPFC). However, very little is known about how direct gaze is processed during live interaction with a real partner. Here, for the first time we used an experimental setup allowing the participant inside an MRI scanner to interact face-to-face with a partner located in the scanner room. Depending on condition, the participant and the partner were instructed either to look at each other in the eyes or to direct their gaze away from the other. As control conditions, participants gazed at their own eyes, reflected in a mirror, or gazed at a picture of the partner's eyes. Results revealed that direct gaze by the partner was associated with activity in areas involved in production and comprehension of language and action, including the IFG, the premotor cortex (PM), and the supplementary motor area (SMA). Activations in these areas were observed regardless of the participant's gaze behavior. In contrast, increased activity in arMPFC, an area involved in inference of other mental states during social interaction and communication, was only observed when the participant reciprocated the partner's direct gaze so as to establish mutual gaze. Psychophysiological interaction (PPI) analysis revealed effective connectivity between the IFG and the arMPFC during mutual gaze. This suggests that, within a larger network concerned with the processing of social gaze, mutual gaze with a real partner is established by an increased coupling between areas involved in the detection of communicative intentions, language, and social interaction.

A neuro-computational model of economic decisions.

Neuronal recordings and lesion studies indicate that key aspects of economic decisions take place in the orbitofrontal cortex (OFC). Previous work identified in this area three groups of neurons encoding the offer value, the chosen value, and the identity of the chosen good. An important and open question is whether and how decisions could emerge from a neural circuit formed by these three populations. Here we adapted a biophysically realistic neural network previously proposed for perceptual decisions (Wang XJ. Neuron 36: 955-968, 2002; Wong KF, Wang XJ. J Neurosci 26: 1314-1328, 2006). The domain of economic decisions is significantly broader than that for which the model was originally designed, yet the model performed remarkably well. The input and output nodes of the network were naturally mapped onto two groups of cells in OFC. Surprisingly, the activity of interneurons in the network closely resembled that of the third group of cells, namely, chosen value cells. The model reproduced several phenomena related to the neuronal origins of choice variability. It also generated testable predictions on the excitatory/inhibitory nature of different neuronal populations and on their connectivity. Some aspects of the empirical data were not reproduced, but simple extensions of the model could overcome these limitations. These results render a biologically credible model for the neuronal mechanisms of economic decisions. They demonstrate that choices could emerge from the activity of cells in the OFC, suggesting that chosen value cells directly participate in the decision process. Importantly, Wang's model provides a platform to investigate the implications of neuroscience results for economic theory.

Subcortical intelligence: caudate volume predicts IQ in healthy adults.

This study examined the association between size of the caudate nuclei and intelligence. Based on the central role of the caudate in learning, as well as neuroimaging studies linking greater caudate volume to better attentional function, verbal ability, and dopamine receptor availability, we hypothesized the existence of a positive association between intelligence and caudate volume in three large independent samples of healthy adults (total N = 517). Regression of IQ onto bilateral caudate volume controlling for age, sex, and total brain volume indicated a significant positive correlation between caudate volume and intelligence, with a comparable magnitude of effect across each of the three samples. No other subcortical structures were independently associated with IQ, suggesting a specific biological link between caudate morphology and intelligence.

Intelligence moderates neural responses to monetary reward and punishment.

The relations between intelligence (IQ) and neural responses to monetary gains and losses were investigated in a simple decision task. In 94 healthy adults, typical responses of striatal blood oxygen level-dependent (BOLD) signal after monetary reward and punishment were weaker for subjects with higher IQ. IQ-moderated differential responses to gains and losses were also found for regions in the medial prefrontal cortex, posterior cingulate cortex, and left inferior frontal cortex. These regions have previously been identified with the subjective utility of monetary outcomes. Analysis of subjects' behavior revealed a correlation between IQ and the extent to which choices were related to experienced decision outcomes in preceding trials. Specifically, higher IQ predicted behavior to be more strongly correlated with an extended period of previously experienced decision outcomes, whereas lower IQ predicted behavior to be correlated exclusively to the most recent decision outcomes. We link these behavioral and imaging findings to a theoretical model capable of describing a role for intelligence during the evaluation of rewards generated by unknown probabilistic processes. Our results demonstrate neural differences in how people of different intelligence respond to experienced monetary rewards and punishments. Our theoretical discussion offers a functional description for how these individual differences may be linked to choice behavior. Together, our results and model support the hypothesis that observed correlations between intelligence and preferences may be rooted in the way decision outcomes are experienced ex post, rather than deriving exclusively from how choices are evaluated ex ante.

Medial prefrontal cortex and striatum mediate the influence of social comparison on the decision process.

We compared private and social decision making to investigate the neural underpinnings of the effect of social comparison on risky choices. We measured brain activity using functional MRI while participants chose between two lotteries: in the private condition, they observed the outcome of the unchosen lottery, and in the social condition, the outcome of the lottery chosen by another person. The striatum, a reward-related brain structure, showed higher activity when participants won more than their counterpart (social gains) compared with winning in isolation and lower activity when they won less than their counterpart (social loss) compared with private loss. The medial prefrontal cortex, implicated in social reasoning, was more activated by social gains than all other events. Sensitivity to social gains influenced both brain activity and behavior during subsequent choices. Specifically, striatal activity associated with social gains predicted medial prefrontal cortex activity during social choices, and experienced social gains induced more risky and competitive behavior in later trials. These results show that interplay between reward and social reasoning networks mediates the influence of social comparison on the decision process.

Working memory and attention in choice.

We study the role of attention and working memory in choices where options are presented sequentially rather than simultaneously. We build a model where a costly attention effort is chosen, which can vary over time. Evidence is accumulated proportionally to this effort and the utility of the reward. Crucially, the evidence accumulated decays over time. Optimal attention allocation maximizes expected utility from final choice; the optimal solution takes the decay into account, so attention is preferentially devoted to later times; but convexity of the flow attention cost prevents it from being concentrated near the end. We test this model with a choice experiment where participants observe sequentially two options. In our data the option presented first is, everything else being equal, significantly less likely to be chosen. This recency effect has a natural explanation with appropriate parameter values in our model of leaky evidence accumulation, where the decline is stronger for the option observed first. Analysis of choice, response time and brain imaging data provide support for the model. Working memory plays an essential role. The recency bias is stronger for participants with weaker performance in working memory tasks. Also activity in parietal areas, coding the stored value in working, declines over time as predicted.

Equality versus self-interest in the brain: differential roles of anterior insula and medial prefrontal cortex.

Everything else being the same, an equal outcome is generally preferred; however, an equitable allocation sometimes is possible only by sacrificing the total amount of resources available to society. Moreover, direct interests may interact with the perception of equality. Here, we have investigated individual preferences, and their neural basis, by employing a task in which an allocation of a fixed amount between the subject and another person (MS condition) or two third parties (TP condition) is randomly determined. The subject can accept or reject the outcome, in the same fashion as the Ultimatum Game: thus an unequal offer may be rejected at the cost of a loss in total amount. Behavioral results show preference for equal outcomes in TP and for equal and advantageous outcomes in MS. An activation of medial prefrontal cortex (mPFC), extending to the anterior middle cingulate cortex (aMCC), was found in MS unequal outcomes, particularly for disadvantageous outcomes and consequent rejections. The anterior insula (AI) was active for unequal outcomes, in both MS and TP. We propose that the equal treatment is a default social norm, and its violation is signaled by the AI, whereas aMCC/mPFC activation, negatively correlated to rejections, reflects the effort to overcome the default rule of equal treatment in favor of a self-advantageous efficiency.

Causes of social reward differences encoded in human brain.

Rewards may be due to skill, effort, and luck, and the social perception of inequality in rewards among individuals may depend on what produced the inequality. Rewards due to skill produce a conflict: higher outcomes of others in this case are considered deserved, and this counters incentives to reduce inequality. However, they also signal superior skill and for this reason induce strong negative affect in those who perform less, which increases the incentive to reduce the inequality. The neurobiological mechanisms underlying evaluation of rewards due to skill, effort, and luck are still unknown. We scanned brain activity of subjects as they perceived monetary rewards caused by skill, effort, or luck. Subjects could subtract from others. Subtraction was larger, everything else being equal, in luck but increased more as the difference in outcomes grew in skill. Similarly, reward-related activation in medial orbitofrontal cortex was more sensitive to the difference in relative outcomes in skill trials. Orbitofrontal activation reflecting comparative reward advantage predicted by how much subjects reduced unfavorable reward inequality later on in the trial. Thus medial orbitofrontal cortex activity reflects the causes of reward and predicts actions that reduce inequality.