Cognitive control moderates early childhood temperament in predicting social behavior in 7‐year‐old children: an ERP study

Behavioral inhibition (BI) is a temperament associated with heightened vigilance and fear of novelty in early childhood, and social reticence and increased risk for anxiety problems later in development. However, not all behaviorally inhibited children develop signs of anxiety. One mechanism that might contribute to the variability in developmental trajectories is the recruitment of cognitive‐control resources. The current study measured N2 activation, an ERP (event‐related potential) associated with cognitive control, and modeled source‐space activation (LORETA; Low Resolution Brain Electromagnetic Tomography) at 7 years of age while children performed a go/no‐go task. Activation was estimated for the entire cortex and then exported for four regions of interest: ventromedial prefrontal cortex (VMPFC), ventrolateral prefrontal cortex (VLPFC), dorsal anterior cingulate cortex (dorsal ACC), and dorsal lateral prefrontal cortex (DLPFC). BI was measured in early childhood (ages 2 and 3 years). Anxiety problems and social reticence were measured at 7 years of age to ascertain stability of temperamental style. Results revealed that BI was associated with increased performance accuracy, longer reaction times, greater (more negative) N2 activation, and higher estimated dorsal ACC and DLPFC activation. Furthermore, early BI was only associated with social reticence at age 7 at higher (more negative) levels of N2 activation or higher estimated dorsal ACC or DLPFC activation. Results are discussed in the context of overcontrolled behavior contributing to social reticence and signs of anxiety in middle childhood.     

Amygdala functional connectivity is reduced after the cold pressor task

The amygdala forms a crucial link between central pain and stress systems. Previous research indicates that psychological stress affects amygdala activity, but it is less clear how painful stressors influence subsequent amygdala functional connectivity. In the present study, we used pulsed arterial spin labeling (PASL) to investigate differences in healthy male adults’ resting-state amygdala functional connectivity following a cold pressor versus a control task, with the stressor and control conditions being conducted on different days. During the period of peak cortisol response to acute stress (approximately 15–30 min after stressor onset), participants were asked to rest for 6 min with their eyes closed during a PASL scanning sequence. The cold pressor task led to reduced resting-state functional connectivity between the amygdalae and orbitofrontal cortex (OFC) and ventromedial prefrontal cortex, and this occurred irrespective of cortisol release. The stressor also induced greater inverse connectivity between the left amygdala and dorsal anterior cingulate cortex (ACC), a brain region implicated in the down-regulation of amygdala responsivity. Furthermore, the degree of poststressor left amygdala decoupling with the lateral OFC varied according to self-reported pain intensity during the cold pressor task. These findings indicate that the cold pressor task alters amygdala interactions with prefrontal and ACC regions 15–30 min after the stressor, and that these altered functional connectivity patterns are related to pain perception rather than cortisol feedback.     

Developmental differences in prefrontal activation during working memory maintenance and manipulation for different memory loads

The ability to keep information active in working memory is one of the cornerstones of cognitive development. Prior studies have demonstrated that regions which are important for working memory performance in adults, such as dorsolateral prefrontal cortex (DLPFC), ventrolateral prefrontal cortex (VLPFC), and superior parietal cortex, become increasingly engaged across school-aged development. The primary goal of the present functional MRI study was to investigate the involvement of these regions in the development of working memory manipulation relative to maintenance functions under different loads. We measured activation in DLPFC, VLPFC, and superior parietal cortex during the delay period of a verbal working memory task in 11-13-year-old children and young adults. We found evidence for age-related behavioral improvements in working memory and functional changes within DLPFC and VLPFC activation patterns. Although activation profiles of DLPFC and VLPFC were similar, group differences were most pronounced for right DLPFC. Consistent with prior studies, right DLPFC showed an interaction between age and condition (i.e. manipulation versus maintenance), specifically at the lower loads. This interaction was characterized by increased activation for manipulation relative to maintenance trials in adults compared to children. In contrast, we did not observe a significant age-dependent load sensitivity. These results suggest that age-related differences in the right DLPFC are specific to working memory manipulation and are not related to task difficulty and/or differences in short-term memory capacity.     

Co-occurring anxiety influences patterns of brain activity in depression

Brain activation associated with anhedonic depression and co-occurring anxious arousal and anxious apprehension was measured by fMRI during performance of an emotion word Stroop task. Consistent with EEG findings, depression was associated with rightward frontal lateralization in the dorsolateral prefrontal cortex (DLPFC), but only when anxious arousal was elevated and anxious apprehension was low. Activity in the right inferior frontal gyrus (IFG) was also reduced for depression under the same conditions. In contrast, depression was associated with more activity in the anterior cingulate cortex (dorsal ACC and rostral ACC) and the bilateral amygdala. Results imply that depression, particularly when accompanied by anxious arousal, may result in a failure to implement top-down processing by appropriate brain regions (left DLPFC, right IFG) due to increased activation in regions associated with responding to emotionally salient information (right DLPFC, amygdala).     

Subliminal food images compromise superior working memory performance in women with restricting anorexia nervosa

Prefrontal cortex (PFC) is dysregulated in women with restricting anorexia nervosa (RAN). It is not known whether appetitive non-conscious stimuli bias cognitive responses in those with RAN. Thirteen women with RAN and 20 healthy controls (HC) completed a dorsolateral PFC (DLPFC) working memory task and an anterior cingulate cortex (ACC) conflict task, while masked subliminal food, aversive and neutral images were presented. During the DLPFC task, accuracy was higher in the RAN compared to the HC group, but superior performance was compromised when subliminal food stimuli were presented: errors positively correlated with self-reported trait anxiety in the RAN group. These effects were not observed in the ACC task. Appetitive activation is intact and anxiogenic in women with RAN, and non-consciously interacts with working memory processes associated with the DLPFC. This interaction mechanism may underlie cognitive inhibition of appetitive processes that are anxiety inducing, in people with AN.     

Brain oscillatory activity associated with task switching and feedback processing

In this study, we sought to dissociate event-related potentials (ERPs) and the oscillatory activity associated with signals indicating feedback about performance (outcome-based behavioral adjustment) and the signals indicating the need to change or maintain a task set (rule-based behavioral adjustment). With this purpose in mind, we noninvasively recorded electroencephalographic signals, using a modified version of the Wisconsin card sorting task, in which feedback processing and task switching could be studied separately. A similar late positive component was observed for the switch and correct feedback signals on the first trials of a series, but feedback-related negativity was observed only for incorrect feedback. Moreover, whereas theta power showed a significant increase after a switch cue and after the first positive feedback of a new series, a selective frontal beta–gamma increase was observed exclusively in the first positive feedback (i.e., after the selection of the new rule). Importantly, for the switch cue, beta–alpha activity was suppressed rather than increased. This clear dissociation between the cue and feedback stimuli in task switching emphasizes the need to accurately study brain oscillatory activity to disentangle the role of different cognitive control processes.     

Dorsolateral prefrontal cortex and Stroop performance: Tackling the lateralization

Neuroscience research has identified the involvement of the dorsolateral prefrontal cortex (DLPFC) in cognitive control. Questions remain, however, about its lateralization correlates during Stroop task performance, an experimental cornerstone on which a large amount of cognitive control research is based. After reviewing the literature, we find that three Stroop variants have been used in an attempt to uncover different aspects of cognitive control related to DLPFC involvement. In sum, rapid and sequential up-regulation of the attentional set seems to be related to the left DLPFC. These attentional adjustments are based on participants’ expectancies regarding the conflicting nature of the upcoming trial, and not on the conflict itself. In contrast, the right DLPFC is associated with an overall up-regulation of the attentional set when attentional conflict is experienced.     

Executive function as a function of sub-clinical psychopathy

Some aspects of executive function are thought to be dysfunctional in psychopathic individuals. We administered a small battery of neuropsychological tests (spatial alternation task, object alternation task, and Porteus Maze) to two samples of college students and obtained a measure of psychopathy via a self-report questionnaire. Psychopathic traits were related to the tests of object alternation and Porteus Maze but not to the spatial alternation task. Our results support the hypothesis of orbitofrontal cortex (OFC) dysfunction with sparing of dorsolateral prefrontal cortex (DLPFC) in psychopathy and provide a downward extension of this theory to sub-clinical levels of psychopathy.     

The effect of motivation on working memory: an fMRI and SEM study

This study investigated the effective connectivity between prefrontal regions of human brain supporting motivational influence on working memory. Functional magnetic resonance imaging (fMRI) and structural equation modeling (SEM) were used to examine the interaction between the lateral orbitofrontal (OFC), medial OFC, and dorsolateral prefrontal (DLPFC) regions in the left and right hemisphere during performance of the verbal 2-back working memory task under two reinforcement conditions. The "low-motivation" condition was not associated with monetary reinforcement, while the "high-motivation" condition involved the probability of winning a certain amount of money. In the "low-motivation" condition, the OFC regions in both hemispheres positively influenced the left DLPFC activity. In the "high-motivation" condition, the connectivity in the network including the right OFC regions and left DLPFC changed from positive to negative, whereas the positive connectivity in the network composed of the left OFC and left DLPFC became slightly enhanced compared with the "low-motivation" condition. However, only the connection between the right lateral OFC and left DLPFC showed a significant condition-dependent change in the strength of influence conveyed through the pathway. This change appears to be the functional correlate of motivational influence on verbal working memory.     

Neurocognitive development of relational reasoning

Relational reasoning is an essential component of fluid intelligence, and is known to have a protracted developmental trajectory. To date, little is known about the neural changes that underlie improvements in reasoning ability over development. In this event‐related functional magnetic resonance imaging (fMRI) study, children aged 8–12 and adults aged 18–25 performed a relational reasoning task adapted from Raven's Progressive Matrices. The task included three levels of relational reasoning demands: REL‐0, REL‐1, and REL‐2. Children exhibited disproportionately lower accuracy than adults on trials that required integration of two relations (REL‐2). Like adults, children engaged lateral prefrontal cortex (PFC) and parietal cortex during task performance; however, they exhibited different time courses and activation profiles, providing insight into their approach to the problems. As in prior studies, adults exhibited increased rostrolateral PFC (RLPFC) activation when relational integration was required (REL‐2 > REL‐1, REL‐0). Children also engaged RLPFC most strongly for REL‐2 problems at early stages of processing, but this differential activation relative to REL‐1 trials was not sustained throughout the trial. These results suggest that the children recruited RLPFC while processing relations, but failed to use it to integrate across two relations. Relational integration is critical for solving a variety of problems, and for appreciating analogies; the current findings suggest that developmental improvements in this function rely on changes in the profile of engagement of RLPFC, as well as dorsolateral PFC and parietal cortex.     

Neural correlates of arithmetic calculation strategies

Recent research into math cognition has identified areas of the brain that are involved in number processing (Dehaene, Piazza, Pinel, & Cohen, 2003) and complex problem solving (Anderson, 2007). Much of this research assumes that participants use a single strategy; yet, behavioral research finds that people use a variety of strategies (LeFevre et al., 1996; Siegler, 1987; Siegler & Lemaire, 1997). In the present study, we examined cortical activation as a function of two different calculation strategies for mentally solving multidigit multiplication problems. The school strategy, equivalent to long multiplication, involves working from right to left. The expert strategy, used by “lightning” mental calculators (Staszewski, 1988), proceeds from left to right. The two strategies require essentially the same calculations, but have different working memory demands (the school strategy incurs greater demands). The school strategy produced significantly greater early activity in areas involved in attentional aspects of number processing (posterior superior parietal lobule, PSPL) and mental representation (posterior parietal cortex, PPC), but not in a numerical magnitude area (horizontal intraparietal sulcus, HIPS) or a semantic memory retrieval area (lateral inferior prefrontal cortex, LIPFC). An ACT-R model of the task successfully predicted BOLD responses in PPC and LIPFC, as well as in PSPL and HIPS.     

Errors without conflict: implications for performance monitoring theories of anterior cingulate cortex

Recent theories of the neural basis of performance monitoring have emphasized a central role for the anterior cingulate cortex (ACC). Replicating an earlier event-related potential (ERP) study, which showed an error feedback negativity that was modeled as having an ACC generator, we used event-related fMRI to investigate whether the ACC would differentiate between correct and incorrect feedback stimuli in a time estimation task. The design controlled for response conflict and frequency and expectancy effects. Although participants in the current study adjusted their performance following error feedback, we did not observe error feedback-evoked ACC activity. In contrast, we did observe ACC activity while the same subjects performed the Stroop task, in which an area of the ACC activated during both conflict and error trials. These findings are inconsistent with previous dipole models of the error feedback negativity, and suggest the ACC may not be involved in the generation of this ERP component. These results question involvement of the ACC in the detection of errors per se when controlling for conflict.     

Inhibition and the right inferior frontal cortex

It is controversial whether different cognitive functions can be mapped to discrete regions of the prefrontal cortex (PFC). The localisationist tradition has associated one cognitive function - inhibition - by turns with dorsolateral prefrontal cortex (DLPFC), inferior frontal cortex (IFC), or orbital frontal cortex (OFC). Inhibition is postulated to be a mechanism by which PFC exerts its effects on subcortical and posterior-cortical regions to implement executive control. We review evidence concerning inhibition of responses and task-sets. Whereas neuroimaging implicates diverse PFC foci, advances in human lesion-mapping support the functional localization of such inhibition to right IFC alone. Future research should investigate the generality of this proposed inhibitory function to other task domains, and its interaction within a wider network.     

Anterior cingulate cortex and conflict detection: An update of theory and data

The dorsal anterior cingulate cortex (ACC) and associated regions of the medial frontal wall have often been hypothesized to play an important role in cognitive control. We have proposed that the ACC's specific role in cognitive control is to detect conflict between simultaneously active, competing representations and to engage the dorsolateral prefrontal cortex (DLPFC) to resolve such conflict. Here we review some of the evidence supporting this theory, from event-related potential (ERP) and fMRI studies. We focus on data obtained from interference tasks, such as the Stroop task, and review the evidence that trial-to-trial changes in control engagement can be understood as driven by conflict detection; the data suggest that levels of activation of the ACC and the DLPFC in such tasks do indeed reflect conflict and control, respectively. We also discuss some discrepant results in the literature that highlight the need for future research.     

Cortical regions activated by the subjective sense of perceptual coherence of environmental sounds: A proposal for a neuroscience of intuition

According to the Oxford English Dictionary, intuition is “the ability to understand or know something immediately, without conscious reasoning.” In other words, people continuously, without conscious attention, recognize patterns in the stream of sensations that impinge upon them. The result is a vague perception of coherence, which subsequently biases thought and behavior accordingly. Within the visual domain, research using paradigms with difficult recognition has suggested that the orbitofrontal cortex (OFC) serves as a fast detector and predictor of potential content that utilizes coarse facets of the input. To investigate whether the OFC is crucial in biasing task-specific processing, and hence subserves intuitive judgments in various modalities, we used a difficult-recognition paradigm in the auditory domain. Participants were presented with short sequences of distorted, nonverbal, environmental sounds and had to perform a sound categorization task. Imaging results revealed rostral medial OFC activation for such auditory intuitive coherence judgments. By means of a conjunction analysis between the present results and those from a previous study on visual intuitive coherence judgments, the rostral medial OFC was shown to be activated via both modalities. We conclude that rostral OFC activation during intuitive coherence judgments subserves the detection of potential content on the basis of only coarse facets of the input.     

The contribution of distinct subregions of the ventromedial frontal cortex to emotion,social behavior,and decision making

Damage to the ventromedial frontal cortex (VMFC) in humans is associated with deficits in decision making. Decision making, however, often happens while people are interacting with others, where it is important to take the social consequences of a course of action into account. It is well known that VMFC lesions also lead to marked alterations in patients’ emotions and ability to interact socially; however, it has not been clear which parts of the VMFC are critical for these changes. Recently, there has been considerable interest in the role of the VMFC in choice behavior during interpersonal exchanges. Here, we highlight recent research that suggests that two areas within or adjacent to the VMFC, the orbitofrontal cortex (OFC) and the anterior cingulate cortex (ACC), may play distinct but complementary roles in mediating normal patterns of emotion and social behavior. Converging lines of evidence from human, macaque, and rat studies now suggest that the OFC may be more specialized for simple emotional responses, such as fear and aggression, through its role in representing primary reinforcement or punishment. By contrast, the ACC may play a distinct role in more complex aspects of emotion, such as social interaction, by virtue of its connections with the discrete parts of the temporal lobe and subcortical structures that control autonomic responses.     

Working memory in the oldest-old: evidence from output serial position curves

In the present study, we examined adult age differences in short-term and working memory performance in middle-aged (45–64 years), young–old (65–74 years), old–old (75–89 years), and oldest–old adults (90 years and over) in the Louisiana Healthy Aging Study. Previous research suggests that measures of working memory are more sensitive to age effects than are simple tests of short-term memory Bopp and Verhaeghen (Journal of Gerontology: Psychological Sciences 60:223–233, 2005), Myerson, Emery, White, and Hale, (Aging, Neuropsychology, and Cognition 10:20–27, 2003). To test this hypothesis, we examined output serial position curves of recall data from three span tasks: forward and backward digit span and size judgment span. Participants’ recall patterns in the size judgment span task revealed that the two oldest groups of adults showed the largest decreases in recall performance across output serial positions, but did not differ significantly from each other. Correlation analyses indicated the strongest negative correlation with age occurred with the size judgment span task. Implications of these findings for understanding strategic processing abilities in late life are discussed.     

The influence of semantic property and grammatical class on semantic selection

Research to-date has not successfully demonstrated consistent neural distinctions for different types of ambiguity or explored the effect of grammatical class on semantic selection. We conducted a relatedness judgment task using event-related functional magnetic resonance imaging (fMRI) to further explore these topics. Participants judged relatedness within word pairs. Consistent and inconsistent conditions were included along with filler items. Imaging results revealed a main effect of ambiguity in the dorsolateral prefrontal cortex (DLPFC) and parietal cortices. A main effect of grammatical class was observed in the parahippocampal and lingual gyri, and a main effect of consistency was found in the DLPFC, ventrolateral prefrontal cortex (VLPFC) and occipital cortices. Interactions among these factors were observed in the cingulate gyrus and motor cortices in addition to the DLPFC. These results suggest that both ambiguity type and grammatical class modulate semantic selection through different neural regions.     

Alertness in young healthy subjects: An fMRI study of brain region interactivity enhanced by a warning signal

An effective connectivity study was carried out on 16 young, healthy subjects performing an alertness task. The objective of this study was to develop and to evaluate a putative network model of alertness by adapting structural equation modeling to fMRI data. This study was designed to evaluate the directed interactivity of an attentional network during intrinsic and phasic alerting tasks. On the basis of theoretical hypotheses, clinical observations, behavioral data and neuroimaging studies, it was hypothesized that neural circuits in the right hemisphere including the dorsolateral prefrontal cortex (DLPFC), anterior cingulate cortex, inferior parietal cortex and the thalamus were involved. The results of this study support the existence of a common network of activated areas with significant path coefficient differences between intrinsic and phasic alertness. Functional interactivity was significantly reinforced during the phasic alertness task and appeared to preferentially involve activity in the DLPFC region, whereas the path coefficients of the model were well-balanced during intrinsic alertness. This study highlights the predominant role of the DLPFC region in maintenance of a state of alertness and in temporal preparation during an alertness task.