PsychoPy2: Experiments in behavior made easy.

PsychoPy is an application for the creation of experiments in behavioral science (psychology, neuroscience, linguistics, etc.) with precise spatial control and timing of stimuli. It now provides a choice of interface; users can write scripts in Python if they choose, while those who prefer to construct experiments graphically can use the new Builder interface. Here we describe the features that have been added over the last 10 years of its development. The most notable addition has been that Builder interface, allowing users to create studies with minimal or no programming, while also allowing the insertion of Python code for maximal flexibility. We also present some of the other new features, including further stimulus options, asynchronous time-stamped hardware polling, and better support for open science and reproducibility. Tens of thousands of users now launch PsychoPy every month, and more than 90 people have contributed to the code. We discuss the current state of the project, as well as plans for the future.

Openness to Experience and Intellect Differentially Predict Creative Achievement in the Arts and Sciences.

The Big Five personality dimension Openness/Intellect is the trait most closely associated with creativity and creative achievement. Little is known, however, regarding the discriminant validity of its two aspects-Openness to Experience (reflecting cognitive engagement with perception, fantasy, aesthetics, and emotions) and Intellect (reflecting cognitive engagement with abstract and semantic information, primarily through reasoning)-in relation to creativity. In four demographically diverse samples totaling 1,035 participants, we investigated the independent predictive validity of Openness and Intellect by assessing the relations among cognitive ability, divergent thinking, personality, and creative achievement across the arts and sciences. We confirmed the hypothesis that whereas Openness predicts creative achievement in the arts, Intellect predicts creative achievement in the sciences. Inclusion of performance measures of general cognitive ability and divergent thinking indicated that the relation of Intellect to scientific creativity may be due at least in part to these abilities. Lastly, we found that Extraversion additionally predicted creative achievement in the arts, independently of Openness. Results are discussed in the context of dual-process theory.

Frontopolar activity and connectivity support dynamic conscious augmentation of creative state.

No ability is more valued in the modern innovation-fueled economy than thinking creatively on demand, and the "thinking cap" capacity to augment state creativity (i.e., to try and succeed at thinking more creatively) is of broad importance for education and a rich mental life. Although brain-based creativity research has focused on static individual differences in trait creativity, less is known about changes in creative state within an individual. How does the brain augment state creativity when creative thinking is required? Can augmented creative state be consciously engaged and disengaged dynamically across time? Using a novel "thin slice" creativity paradigm in 55 fMRI participants performing verb-generation, we successfully cued large, conscious, short-duration increases in state creativity, indexed quantitatively by a measure of semantic distance derived via latent semantic analysis. A region of left frontopolar cortex, previously associated with creative integration of semantic information, exhibited increased activity and functional connectivity to anterior cingulate gyrus and right frontopolar cortex during cued augmentation of state creativity. Individual differences in the extent of increased activity in this region predicted individual differences in the extent to which participants were able to successfully augment state creative performance after accounting for trait creativity and intelligence.

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.

A gene-brain-cognition pathway: prefrontal activity mediates the effect of COMT on cognitive control and IQ.

A core thesis of cognitive neurogenetic research is that genetic effects on cognitive ability are mediated by specific neural functions, however, demonstrating neural mediation has proved elusive. Pairwise relationships between genetic variation and brain function have yielded heterogeneous findings to date. This heterogeneity indicates that a multiple mediator modeling approach may be useful to account for complex relationships involving function at multiple brain regions. This is relevant not only for characterizing healthy cognition but for modeling the complex neural pathways by which disease-related genetic effects are transmitted to disordered cognitive phenotypes in psychiatric illness. Here, in 160 genotyped functional magnetic resonance imaging participants, we used a multiple mediator model to test a gene-brain-cognition pathway by which activity in 4 prefrontal brain regions mediates the effects of catechol-O-methyltransferase (COMT) gene on cognitive control and IQ. Results provide evidence for gene-brain-cognition mediation and help delineate a pathway by which gene expression contributes to intelligence.

Meditation experience is associated with differences in default mode network activity and connectivity.

Many philosophical and contemplative traditions teach that "living in the moment" increases happiness. However, the default mode of humans appears to be that of mind-wandering, which correlates with unhappiness, and with activation in a network of brain areas associated with self-referential processing. We investigated brain activity in experienced meditators and matched meditation-naive controls as they performed several different meditations (Concentration, Loving-Kindness, Choiceless Awareness). We found that the main nodes of the default-mode network (medial prefrontal and posterior cingulate cortices) were relatively deactivated in experienced meditators across all meditation types. Furthermore, functional connectivity analysis revealed stronger coupling in experienced meditators between the posterior cingulate, dorsal anterior cingulate, and dorsolateral prefrontal cortices (regions previously implicated in self-monitoring and cognitive control), both at baseline and during meditation. Our findings demonstrate differences in the default-mode network that are consistent with decreased mind-wandering. As such, these provide a unique understanding of possible neural mechanisms of meditation.

Openness to experience, intellect, and cognitive ability.

An instrument designed to separate 2 midlevel traits within each of the Big Five (the Big Five Aspect Scales [BFAS]) was used to clarify the relation of personality to cognitive ability. The BFAS measures Openness to Experience and Intellect as separate (although related) traits, and refers to the broader Big Five trait as Openness/Intellect. In 2 samples (N = 125 and 189), Intellect was independently associated with general intelligence (g) and with verbal and nonverbal intelligence about equally. Openness was independently associated only with verbal intelligence. Implications of these findings are discussed for the empirical and conceptual relations of intelligence to personality and for the mechanisms potentially underlying both Openness/Intellect and cognitive ability.

Thin slices of creativity: using single-word utterances to assess creative cognition.

We investigated the hypothesis that individual differences in creative cognition can be manifest even in brief responses, such as single-word utterances. Participants (n = 193) were instructed to say a verb upon seeing a noun displayed on a computer screen and were cued to respond creatively to half of the nouns. For every noun-verb pair (72 pairs per subject), we assessed the semantic distance between the noun and the verb, using latent semantic analysis (LSA). Semantic distance was higher in the cued ("creative") condition than the uncued condition, within subjects. Critically, between subjects, semantic distance in the cued condition had a strong relationship to a creativity factor derived from a battery of verbal, nonverbal, and achievement-based creativity measures (ß= .50), and this relation remained when controlling for intelligence and personality. The data show that creative cognition can be assessed reliably and validly from such thin slices of behavior.

Real-time fMRI links subjective experience with brain activity during focused attention.

Recent advances in brain imaging have improved the measure of neural processes related to perceptual, cognitive and affective functions, yet the relation between brain activity and subjective experience remains poorly characterized. In part, it is a challenge to obtain reliable accounts of participant's experience in such studies. Here we addressed this limitation by utilizing experienced meditators who are expert in introspection. We tested a novel method to link objective and subjective data, using real-time fMRI (rt-fMRI) to provide participants with feedback of their own brain activity during an ongoing task. We provided real-time feedback during a focused attention task from the posterior cingulate cortex, a hub of the default mode network shown to be activated during mind-wandering and deactivated during meditation. In a first experiment, both meditators and non-meditators reported significant correspondence between the feedback graph and their subjective experience of focused attention and mind-wandering. When instructed to volitionally decrease the feedback graph, meditators, but not non-meditators, showed significant deactivation of the posterior cingulate cortex. We were able to replicate these results in a separate group of meditators using a novel step-wise rt-fMRI discovery protocol in which participants were not provided with prior knowledge of the expected relationship between their experience and the feedback graph (i.e., focused attention versus mind-wandering). These findings support the feasibility of using rt-fMRI to link objective measures of brain activity with reports of ongoing subjective experience in cognitive neuroscience research, and demonstrate the generalization of expertise in introspective awareness to novel contexts.

Neural correlates of the essence of conscious conflict: fMRI of sustaining incompatible intentions.

The study of intrapsychic conflict has long been central to many key theories about the control of behavior. More recently, by focusing on the nature of conflicting processes in the brain, investigators have revealed great insights about controlled versus automatic processes and the nature of self-control. Despite these advances, many theories of cognitive control or self-control remain agnostic about the function of subjective awareness (i.e., basic consciousness). Why people consciously experience some conflicts in the nervous system but not others remains a mystery. One hypothesis is that people become conscious only of conflicts involving competition for the control of skeletal muscle. To test one aspect of this larger hypothesis, in the present study, 14 participants were trained to introspect the feeling of conflict (the urge to make an error during a Stroop color-word interference task) and then were asked to introspect in the same way while sustaining simple compatible and incompatible intentions during fMRI scanning (to move a finger left or right). As predicted, merely sustaining incompatible skeletomotor intentions prior to their execution produced stronger systematic changes in subjective experience than sustaining compatible intentions, as indicated by self-report ratings obtained in the scanner. Similar ratings held for a modified Stroop-like task when contrasting incompatible versus compatible trials also during fMRI scanning. We use subjective ratings as the basis of parametric analyses of fMRI data, focusing a priori on the brain regions involved in action-related urges (e.g., parietal cortex) and cognitive control (e.g., dorsal anterior cingulate cortex, lateral PFC). The results showed that subjective conflict from sustaining incompatible intentions was consistently related to activity in the left post-central gyrus.

Interaction of COMT val158met and externalizing behavior: relation to prefrontal brain activity and behavioral performance.

A promising approach in neuroimaging studies aimed at understanding effects of single genetic variants on behavior is the study of gene-trait interactions. Variation in the catechol-O-methyl-transferase gene (COMT) is associated with the regulation of dopamine levels in the prefrontal cortex and with cognitive functioning. Given the involvement of dopaminergic neurotransmission in externalizing behavior, a trait characterized by impulsivity and aggression, especially in men, externalizing (as a trait) may index a set of genetic, environmental, and neural characteristics pertinent to understanding phenotypic effects of genetic variation in the COMT gene. In the current study, we used a gene-trait approach to investigate effects of the COMT val(158)met polymorphism and externalizing on brain activity during moments involving low or high demands on cognitive control. In 104 male participants, interference-related activation depended conjointly on externalizing and val(158)met: stronger activation in the dorsal anterior cingulate and lateral prefrontal cortex was found for val/val individuals with high trait externalizing while stronger activation in cingulate motor areas and sensorimotor precuneus was found for met/met individuals with low externalizing. Our results suggest that the val/val genotype, coupled with high levels of trait externalizing, lowers the efficiency of stimulus conflict resolution, whereas the met/met genotype, coupled with low levels of externalizing, lowers the efficiency of response selection.

Connecting long distance: semantic distance in analogical reasoning modulates frontopolar cortex activity.

Solving problems often requires seeing new connections between concepts or events that seemed unrelated at first. Innovative solutions of this kind depend on analogical reasoning, a relational reasoning process that involves mapping similarities between concepts. Brain-based evidence has implicated the frontal pole of the brain as important for analogical mapping. Separately, cognitive research has identified semantic distance as a key characteristic of the kind of analogical mapping that can support innovation (i.e., identifying similarities across greater semantic distance reveals connections that support more innovative solutions and models). However, the neural substrates of semantically distant analogical mapping are not well understood. Here, we used functional magnetic resonance imaging (fMRI) to measure brain activity during an analogical reasoning task, in which we parametrically varied the semantic distance between the items in the analogies. Semantic distance was derived quantitatively from latent semantic analysis. Across 23 participants, activity in an a priori region of interest (ROI) in left frontopolar cortex covaried parametrically with increasing semantic distance, even after removing effects of task difficulty. This ROI was centered on a functional peak that we previously associated with analogical mapping. To our knowledge, these data represent a first empirical characterization of how the brain mediates semantically distant analogical mapping.

Testing predictions from personality neuroscience. Brain structure and the big five.

We used a new theory of the biological basis of the Big Five personality traits to generate hypotheses about the association of each trait with the volume of different brain regions. Controlling for age, sex, and whole-brain volume, results from structural magnetic resonance imaging of 116 healthy adults supported our hypotheses for four of the five traits: Extraversion, Neuroticism, Agreeableness, and Conscientiousness. Extraversion covaried with volume of medial orbitofrontal cortex, a brain region involved in processing reward information. Neuroticism covaried with volume of brain regions associated with threat, punishment, and negative affect. Agreeableness covaried with volume in regions that process information about the intentions and mental states of other individuals. Conscientiousness covaried with volume in lateral prefrontal cortex, a region involved in planning and the voluntary control of behavior. These findings support our biologically based, explanatory model of the Big Five and demonstrate the potential of personality neuroscience (i.e., the systematic study of individual differences in personality using neuroscience methods) as a discipline.

Multiple bases of human intelligence revealed by cortical thickness and neural activation.

We hypothesized that individual differences in intelligence (Spearman's g) are supported by multiple brain regions, and in particular that fluid (gF) and crystallized (gC) components of intelligence are related to brain function and structure with a distinct profile of association across brain regions. In 225 healthy young adults scanned with structural and functional magnetic resonance imaging sequences, regions of interest (ROIs) were defined on the basis of a correlation between g and either brain structure or brain function. In these ROIs, gC was more strongly related to structure (cortical thickness) than function, whereas gF was more strongly related to function (blood oxygenation level-dependent signal during reasoning) than structure. We further validated this finding by generating a neurometric prediction model of intelligence quotient (IQ) that explained 50% of variance in IQ in an independent sample. The data compel a nuanced view of the neurobiology of intelligence, providing the most persuasive evidence to date for theories emphasizing multiple distributed brain regions differing in function.

Individual differences in delay discounting: relation to intelligence, working memory, and anterior prefrontal cortex.

Lower delay discounting (better self-control) is linked to higher intelligence, but the basis of this relation is uncertain. To investigate the potential role of working memory (WM) processes, we assessed delay discounting, intelligence (g), WM (span tasks, 3-back task), and WM-related neural activity (using functional magnetic resonance imaging) in 103 healthy adults. Delay discounting was negatively correlated with g and WM. WM explained no variance in delay discounting beyond that explained by g, which suggests that processes through which WM relates to delay discounting are shared by g. WM-related neural activity in left anterior prefrontal cortex (Brodmann's area 10) covaried with g, r= .26, and delay discounting, r=-.40, and partially mediated the relation between g and delay discounting. Overall, the results suggest that delay discounting is associated with intelligence in part because of processes instantiated in anterior prefrontal cortex, a region known to support the integration of diverse information.

Neural mechanisms of general fluid intelligence.

We used an individual-differences approach to test whether general fluid intelligence (gF) is mediated by brain regions that support attentional (executive) control, including subregions of the prefrontal cortex. Forty-eight participants first completed a standard measure of gF (Raven's Advanced Progressive Matrices). They then performed verbal and nonverbal versions of a challenging working-memory task (three-back) while their brain activity was measured using functional magnetic resonance imaging (fMRI). Trials within the three-back task varied greatly in the demand for attentional control because of differences in trial-to-trial interference. On high-interference trials specifically, participants with higher gF were more accurate and had greater event-related neural activity in several brain regions. Multiple regression analyses indicated that lateral prefrontal and parietal regions may mediate the relation between ability (gF) and performance (accuracy despite interference), providing constraints on the neural mechanisms that support gF.

Individual differences in amygdala activity predict response speed during working memory.

The human amygdala has classically been viewed as a brain structure primarily related to emotions and dissociated from higher cognition. We report here findings suggesting that the human amygdala also has a role in supporting working memory (WM), a canonical higher cognitive function. In a first functional magnetic resonance imaging (fMRI) study (n = 53), individual differences in amygdala activity predicted behavioral performance in a 3-back WM task. Specifically, higher event-related amygdala amplitude predicted faster response time (RT; r = -0.64), with no loss of accuracy. This relationship was not contingent on mood state, task content, or personality variables. In a second fMRI study (n = 21), we replicated the key finding (r = -0.47) and further showed that the correlation between the amygdala and faster RT was specific to a high working memory load condition (3-back) compared with a low working memory load condition (1-back). These results support models of amygdala function that can account for its involvement not only in emotion but also higher cognition.

A role for the human amygdala in higher cognition.

Considerable evidence suggests that the human amygdala plays an important role in higher cognitive functions in addition to its well-known role in emotional processing. In this article we review representative evidence showing the involvement of the human amygdala in long-term memory, working memory and attention. Results are discussed in terms of their relevance to current theories of amygdala function that can integrate its cognitive and emotional functions in a comprehensive framework.

Sustained neural activity associated with cognitive control during temporally extended decision making.

Decision making has both cognitive and affective components, but previous neuroimaging studies in this domain predominantly have focused on affect and reward. The current study examined a decision-making paradigm that placed strong demands on cognitive control processes by making reward payoffs contingent upon decision-making history. Payoffs were maximized by choosing the option that, paradoxically, was associated with a lower payoff on the immediate trial. Temporal integration requirements were manipulated by varying, across conditions, the window of previous trials over which the reward function was calculated. The cognitive demands of the task were hypothesized to engage neural systems responsible for integrating and actively maintaining actions and outcomes over time and the top-down biasing of response selection. Brain activation was monitored with functional magnetic resonance imaging (fMRI) using a mixed-blocked and event-related design to extract both transient and sustained neural responses. A network of brain regions commonly associated with cognitive control functions, including bilateral prefrontal cortex (PFC), bilateral parietal cortex, and medial frontal cortex, showed selectively sustained activation during the task. Increasing temporal integration demands led to a shift from transient to sustained activity in additional regions, including right hemisphere dorsolateral and frontopolar PFC. These results demonstrate the contribution of cognitive control mechanisms to temporally extended decision-making paradigms and highlight the benefits of decomposing activation responses into sustained and transient components.