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Cameron S. Carter
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Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2020) 32 (11): 2117–2130.
Published: 01 November 2020
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Prior studies demonstrated that neural oscillations are enhanced during working memory (WM) maintenance and that this activity can predict behavioral performance in healthy individuals. However, it is unclear whether the relationship holds for people with WM deficits. People with schizophrenia have marked WM deficits, and such deficits are most prominent when patients are required to process relationships between items, such as temporal order. Here, we used EEG to compare the relationship between oscillatory activity and WM performance in patients and controls. EEG was recorded as participants performed tasks requiring maintenance of complex objects (“Item”) or the temporal order of objects (“Order”). In addition to testing for group differences, we examined individual differences in EEG power and WM performance across groups. Behavioral results demonstrated that patients showed impaired performance on both Item and Order trials. EEG analyses revealed that patients showed an overall reduction in alpha power, but the relationship between alpha activity and performance was preserved. In contrast, patients showed a reduction in theta power specific to Order trials, and theta power could predict performance on Order trials in controls, but not in patients. These findings demonstrate that WM impairments in patients may reflect two different processes: a general deficit in alpha oscillations and a specific deficit in theta oscillations when temporal order information must be maintained. At a broader level, the results highlight the value of characterizing brain–behavior relationships, by demonstrating that the relationship between neural oscillations and WM performance can be fundamentally disrupted in those with WM deficits.
Journal Articles
Subthalamic Nucleus Activation Occurs Early during Stopping and Is Associated with Trait Impulsivity
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2019) 31 (4): 510–521.
Published: 01 April 2019
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The subthalamic nucleus (STN) is thought to be a central regulator of behavioral inhibition, which is thought to be a major determinant of impulsivity. Thus, it would be reasonable to hypothesize that STN function is related to impulsivity. However, it has been difficult to test this hypothesis because of the challenges in noninvasively and accurately measuring this structure's signal in humans. We utilized a novel approach for STN signal localization that entails identifying this structure directly on fMRI images for each individual participant in native space. Using this approach, we measured STN responses during the stop signal task in a sample of healthy adult participants. We confirmed that the STN exhibited selective activation during “Stop” trials. Furthermore, the magnitude of STN activation during successful Stop trials inversely correlated with individual differences in trait impulsivity as measured by a personality inventory. Time course analysis revealed that early STN activation differentiated successful from unsuccessful Stop trials, and individual differences in the magnitude of STN activation inversely correlated with stop signal RT, an estimate of time required to stop. These results are consistent with the STN playing a central role in inhibition and related behavioral proclivities, with implications for both normal range function and clinical syndromes of inhibitory dyscontrol. Moreover, the methods utilized in this study for measuring STN fMRI signal in humans may be gainfully applied in future studies to further our understanding of the role of the STN in regulating behavior and neuropsychiatric conditions.
Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2015) 27 (12): 2309–2323.
Published: 01 December 2015
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The establishment of reference is essential to language comprehension. The goal of this study was to examine listeners' sensitivity to referential ambiguity as a function of individual variation in attention, working memory capacity, and verbal ability. Participants listened to stories in which two entities were introduced that were either very similar (e.g., two oaks) or less similar (e.g., one oak and one elm). The manipulation rendered an anaphor in a subsequent sentence (e.g., oak) ambiguous or unambiguous. EEG was recorded as listeners comprehended the story, after which participants completed tasks to assess working memory, verbal ability, and the ability to use context in task performance. Power in the alpha and theta frequency bands when listeners received critical information about the discourse entities (e.g., oaks) was used to index attention and the involvement of the working memory system in processing the entities. These measures were then used to predict an ERP component that is sensitive to referential ambiguity, the Nref, which was recorded when listeners received the anaphor. Nref amplitude at the anaphor was predicted by alpha power during the earlier critical sentence: Individuals with increased alpha power in ambiguous compared with unambiguous stories were less sensitive to the anaphor's ambiguity. Verbal ability was also predictive of greater sensitivity to referential ambiguity. Finally, increased theta power in the ambiguous compared with unambiguous condition was associated with higher working-memory span. These results highlight the role of attention and working memory in referential processing during listening comprehension.
Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2015) 27 (6): 1238–1248.
Published: 01 June 2015
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The ability to proactively control motor responses, particularly to overcome overlearned or automatic actions, is an essential prerequisite for adaptive, goal-oriented behavior. The substantia nigra (SN), an element of the BG, has figured prominently in current models of response selection. However, because of its small size and proximity to functionally distinct subcortical structures, it has been challenging to test the SN's involvement in response selection using conventional in vivo functional neuroimaging approaches. We developed a new fMRI localization method for directly distinguishing, on echo-planar images, the SN BOLD signal from that of neighboring structures, including the subthalamic nucleus (STN). Using this method, we tested the hypothesis that the SN supports the proactive control of response selection. We acquired high-resolution EPI volumes at 3 T from 16 healthy participants while they completed the Preparing to Overcome Prepotency task of proactive control. There was significantly elevated delay period signal selectively during high- compared with low-control trials in the SN. The STN did not show delay period activity in either condition. SN delay period signal was significantly inversely associated with task performance RTs across participants. These results suggest that our method offers a novel means for measuring SN BOLD responses, provides unique evidence of SN involvement in cognitive control in humans, and suggests a novel mechanism for proactive response selection.
Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2011) 23 (9): 2211–2221.
Published: 01 September 2011
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Human behavior involves monitoring and adjusting performance to meet established goals. Performance-monitoring systems that act by detecting conflict in stimulus and response processing have been hypothesized to influence cortical control systems to adjust and improve performance. Here we used fMRI to investigate the neural mechanisms of conflict monitoring and resolution during voluntary spatial attention. We tested the hypothesis that the ACC would be sensitive to conflict during attentional orienting and influence activity in the frontoparietal attentional control network that selectively modulates visual information processing. We found that activity in ACC increased monotonically with increasing attentional conflict. This increased conflict detection activity was correlated with both increased activity in the attentional control network and improved speed and accuracy from one trial to the next. These results establish a long hypothesized interaction between conflict detection systems and neural systems supporting voluntary control of visual attention.
Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2011) 23 (4): 923–935.
Published: 01 April 2011
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Mechanisms by which the brain monitors and modulates performance are an important focus of recent research. The conflict-monitoring hypothesis posits that the ACC detects conflict between competing response pathways which, in turn, signals for enhanced control. The N2, an ERP component that has been localized to ACC, has been observed after high conflict stimuli. As a candidate index of the conflict signal, the N2 would be expected to be sensitive to the degree of response conflict present, a factor that depends on both the features of external stimuli and the internal control state. In the present study, we sought to explore the relationship between N2 amplitude and these variables through use of a modified Eriksen flankers task in which target–distracter compatibility was parametrically varied. We hypothesized that greater target–distracter incompatibility would result in higher levels of response conflict, as indexed by both behavior and the N2 component. Consistent with this prediction, there were parametric degradations in behavioral performance and increases in N2 amplitudes with increasing incompatibility. Further, increasingly incompatible stimuli led to the predicted parametric increases in control on subsequent incompatible trials as evidenced by enhanced performance and reduced N2 amplitudes. These findings suggest that the N2 component and associated behavioral performance are finely sensitive to the degree of response conflict present and to the control adjustments that result from modulations in conflict.
Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2008) 20 (11): 1952–1965.
Published: 01 November 2008
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People are capable, at will, of trading speed for accuracy when performing a task; they can focus on performing accurately at the cost of being slow, or emphasize speed at the cost of decreased accuracy. Here, we used functional magnetic resonance imaging to investigate the neural correlates of this ability. We show increased baseline activity during speed emphasis in a network of areas related to response preparation and execution, including the premotor areas of the frontal lobe, the basal ganglia, the thalamus, and the dorsolateral prefrontal and left parietal cortices. Furthermore, speed emphasis was associated with reduced transient response-related activation in several of these structures, suggesting that because of the greater baseline activity under speed emphasis, less activation is needed in these structures to reach response threshold, consistent with the assumptions of several computational theories. Moreover, we identify the dorsolateral prefrontal cortex as providing the top-down control signal that increases this baseline activity.
Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2008) 20 (4): 682–693.
Published: 01 April 2008
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Brain imaging studies suggest that the rostrolateral prefrontal cortex (RLPFC), is involved in relational reasoning. Functional magnetic resonance imaging (fMRI) studies involving Raven's Progressive Matrices or verbal propositional analogies indicate that the RLPFC is engaged by tasks that require integration across multiple relational structures. Several studies have shown that the RLPFC is more active when people must evaluate an analogy (e.g., Is shoe to foot as glove is to hand?) than when they must simply evaluate two individual semantic relationships, consistent with the hypothesis that this region is important for relational integration. The current fMRI investigation further explores the role of the RLPFC in reasoning and relational integration by comparing RLPFC activation across four different propositional analogy conditions. Each of the four conditions required either relation completion (e.g., Shoe is to foot as glove is to WHAT? → “hand”) or relation comparison (e.g., Is shoe to foot as glove is to hand? → “yes”). The RLPFC was engaged more strongly by the comparison subtask relative to completion, suggesting that the RLPFC is particularly involved in comparing relational structures.
Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2004) 16 (5): 786–793.
Published: 01 June 2004
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Several previous studies have compared the blood oxygen level-dependent (BOLD) hemodynamic response (HDR) in healthy elderly subjects to the HDR in young subjects. Some studies have found a relative decreased amplitude in the elderly in the visual cortex, whereas other studies have found the elderly HDR amplitude in the visual cortex to be nearly identical to that in young subjects. A possible explanation for the different findings is that the peak voxel HDR is similar between the groups, but that the HDR in the group-averaged region-of-interest (ROI) is “washed out” by the inclusion of less significant voxels (due to a smaller extent of activation in the elderly) or by the inclusion of negative-peaking voxels. We tested this hypothesis using event-related functional magnetic resonance imaging (fMRI). While undergoing fMRI, subjects performed a simple visual and motor task, pressing with their index fingers in response to visual presentation of the word tap. Data from 18 subjects, 8 young and 10 elderly, were analyzed. For each subject, a visual and a motor ROI was selected by choosing the most significant positive voxels within the anatomically defined ROI. This individual subject approach excluded both low-significance and negative-peaking voxels. Similar peaks were found for the elderly and the young subjects in both motor and visual regions and a more sustained BOLD response was found for the elderly in both regions. Additionally, as predicted, a greater percentage of voxels with a negative HDR was found for the elderly in the visual region; this finding was also replicated in our reanalysis of an independent fMRI and aging study from the fMRI Data Center. Functional neuroimaging observations of negative HDRs in visual areas have been interpreted as the effect of unconstrained processing during rest. Our results suggest that the elderly may have more unconstrained visual processing during the rest condition in the scanner. The observation that the group differences in the BOLD response are sensitive to voxel selection (e.g., inclusion of low-significance and/or negative voxels) underscores the importance of ROI selection criteria in the interpretation of fMRI studies using elderly populations.
Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2004) 16 (4): 637–653.
Published: 01 May 2004
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This research tests a model of the computational role of three cortical regions in tasks like algebra equation solving. The model assumes that there is a left parietal region-of-interest (ROI) where the problem expression is represented and transformed, a left prefrontal ROI where information for solving the task is retrieved, and a motor ROI where hand movements to produce the answer are programmed. A functional magnetic resonance imaging (fMRI) study of an abstract symbolmanipulation task was performed to articulate the roles of these three regions. Participants learned to associate words with instructions for transforming strings of letters. The study manipulated the need to retrieve these instructions, the need to transform the strings, and whether there was a delay between calculation of the answer and the output of the answer. As predicted, the left parietal ROI mainly reflected the need for a transformation and the left prefrontal ROI the need for retrieval. Homologous right ROIs showed similar but weaker responses. Neither the prefrontal nor the parietal ROIs responded to delay, but the motor ROI did respond to delay, implying motor rehearsal over the delay. Except for the motor ROI, these patterns of activity did not vary with response hand. In an ACT-R model, it was shown that the activity of an imaginal buffer predicted the blood oxygen level-dependent (BOLD) response of the parietal ROI, the activity of a retrieval buffer predicted the response of the prefrontal ROI, and the activity of a manual buffer predicted the response of the motor ROI.
Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2002) 14 (4): 593–602.
Published: 15 May 2002
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The anterior cingulate cortex (ACC) has been shown to respond to conflict between simultaneously active, incompatible response tendencies. This area is active during high-conflict correct trials and also when participants make errors. Here, we use the temporal resolution of high-density event-related potentials (ERPs) in combination with source localization to investigate the timing of ACC activity during conflict and error detection. We predicted that the same area of the ACC is active prior to high-conflict correct responses and following erroneous responses. Dipole modeling supported this prediction: The frontocentral N2, occurring prior to the response on correct conflict trials, and the ERN, occurring immediately following error responses, could both be modeled as having a generator in the caudal ACC, suggesting the same process to underlie both peaks. Modeling further suggested that the rostral area of the ACC was also active following errors, but later in time, contributing to the error positivity ( P E ), and peaking at 200–250 msec following the ERN peak. Despite the inherent limitations of source localization, these data may begin to shed light on the timing of action-monitoring processes. First, the time course of caudal ACC activity follows the time course as predicted by the conflict theory of this region. Second, caudal ACC activity might be temporally dissociated from rostral ACC activity during error trials, which possibly reflects a separate, affective component of the evaluative functions of the ACC.
Journal Articles
Publisher: Journals Gateway
Journal of Cognitive Neuroscience (2000) 12 (6): 977–987.
Published: 01 November 2000
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Event-related fMRI was used to dissociate the neural systems involved in category learning with and without awareness. Ten subjects performed a speeded response category learning task. Functional MR images were acquired during both explicit and implicit learning conditions. Behavioral data showed evidence of learning in both conditions. Functional imaging data showed different activation patterns in implicit and explicit trials. Decreased activation in extrastriate region V3 was found with implicit learning, and increased activation in V3, the medial temporal lobe, and frontal regions were found with explicit learning. These results support the theory that implicit and explicit learning utilize dissociable neural systems. Moreover, in both the implicit and explicit conditions a similar pattern of decreased activation was found in parietal regions. This commonality suggests that these dissociable systems also operate in parallel.