Helge Gebhardt

Relationship between regional hemodynamic activity and simultaneously recorded EEG-theta associated with mental arithmetic-induced workload

Theta increases with workload and is associated with numerous processes including working memory, problem solving, encoding, or self monitoring. These processes, in turn, involve numerous structures of the brain. However, the relationship between regional brain activity and the occurrence of theta remains unclear. In the present study, simultaneous EEG‐fMRI recordings were used to investigate the functional topography of theta. EEG‐theta was enhanced by mental arithmetic‐induced workload. For the EEG‐constrained fMRI analysis, theta‐reference time‐series were extracted from the EEG, reflecting the strength of theta occurrence during the time course of the experiment. Theta occurrence was mainly associated with activation of the insular cortex, hippocampus, superior temporal areas, cingulate cortex, superior parietal, and frontal areas. Though observation of temporal and insular activation is in accord with the theory that theta specifically reflects encoding processes, the involvement of several other brain regions implies that surface‐recorded theta represents comprehensive functional brain states rather than specific processes in the brain. The results provide further evidence for the concept that emergent theta band oscillations represent dynamic functional binding of widely distributed cortical assemblies, essential for cognitive processing. This binding process may form the source of surface‐recorded EEG theta. Hum Brain Mapp, 2006.

Acquisition of typical EEG waveforms during fMRI: SSVEP, LRP, and frontal theta

Recent work has demonstrated the feasibility of simultaneous electroencephalography (EEG) and functional magnetic resonance imaging (fMRI). Virtually no systematic comparisons between EEG recorded inside and outside the MR scanner have been conducted, and it is unknown if different kinds of frequency mix, topography, and domain-specific processing are uniformly recordable within the scanner environment. The aim of the study was to investigate several typical EEG waveforms in the same subjects inside the magnet during fMRI and outside the MR examination room. We examined whether uniform artifact subtraction allows the extraction of these different EEG waveforms inside the scanner during EPI scanning to the same extent as outside the scanner. Three well-established experiments were conducted, eliciting steady state visual evoked potentials (SSVEP), lateralized readiness potentials (LRP), and frontal theta enhancement induced by mental addition. All waveforms could be extracted from the EEG recorded during fMRI. Substantially no differences in these waveforms of interest were found between gradient-switching and intermediate epochs during fMRI (only the SSVEP-experiment was designed for a comparison of gradient-with intermediate epochs), or between waveforms recorded inside the scanner during EPI scanning and outside the MR examination room (all experiments). However, non-specific amplitude differences were found between inside and outside recorded EEG at lateral electrodes, which were not in any interaction with the effects of interest. The source of these differences requires further exploration. The high concordance of activation patterns with published results demonstrates that EPI-images could be acquired during EEG recording without significant distortion.

Functional correlates of distractor suppression during spatial working memory encoding.

Executive working memory operations are related to prefrontal regions in the healthy brain. Moreover, neuroimaging data provide evidence for a functional dissociation of ventrolateral and dorsolateral prefrontal cortex. Most authors either suggest a modality-specific or a function-specific prefrontal cortex organization. In the present study we particularly aimed at the identification of different prefrontal cerebral areas that are involved in executive inhibitory processes during spatial working memory encoding. In an fMRI study (functional magnetic resonance imaging) we examined the neural correlates of spatial working memory processing by varying the amount of executive demands of the task. Twenty healthy volunteers performed the Corsi Block-Tapping test (CBT) during fMRI. The CBT requires the storage and reproduction of spatial target sequences. In a second condition, we presented an adapted version of the Block-Suppression-Test (BST). The BST is based on the original CBT but additionally requires the active suppression of visual distraction within the target sequences. In comparison to the CBT performance, particularly the left dorsolateral prefrontal cortex (BA 9) showed more activity during the BST condition. Our results show that the left dorsolateral prefrontal cortex plays a crucial role for executive controlled inhibition of spatial distraction. Furthermore, our findings are in line with the processing model of a functional dorsolateral-ventrolateral prefrontal cortex organization.

Hippocampal involvement in working memory encoding of changing locations: An fMRI study

Whereas the role of the hippocampus for spatial learning and long-term memory is largely undisputed, there is less evidence for a participation of hippocampal structures in spatial working memory operations. In an fMRI study (functional magnetic resonance imaging), we therefore examined the role of the hippocampus during spatial working memory performance. Nineteen healthy volunteers performed a modified version of the Corsi Block-Tapping test (CBT) during fMRI. The CBT is a neuropsychological instrument that is widely used in clinical settings. It requires the storage and subsequent reproduction of spatial target sequences. Brain activity during CBT performance has rarely been examined in the past, at least not the hemodynamic correlates. In a baseline condition, participants processed a number of subsequently presented targets as in the CBT condition. The only difference was that targets did not change their location. As compared to baseline activity, the right hippocampus showed more activation during the CBT condition. In addition, whole-brain analysis showed working memory related frontal and parietal brain activation. The results indicate that hippocampal structures contribute to serial working memory encoding of spatial locations in the human brain.

The impact of age on load-related dorsolateral prefrontal cortex activation

Healthy aging is accompanied by working memory-related functional cerebral changes. Depending on performance accuracy and the level of working memory demands, older adults show task-related patterns of either increased or decreased activation compared to younger adults. Controversies remain concerning the interpretation of these changes and whether they already manifest in earlier decades of life. To address these issues, functional magnetic resonance imaging (fMRI) was used to examine brain activation during spatial working memory retrieval in 45 healthy individuals between 20 and 68 years of age. Participants performed a modified version of the Corsi Block-Tapping test (CBT). The CBT requires the storage and subsequent reproduction of spatial target sequences and allows modulating working memory load by a variation of sequence length. Results revealed that activation intensity at the lowest CBT load level increased with increasing age and positively correlated with the number of errors. At higher CBT load levels, activation intensity decreased with increasing age together with a disproportional accuracy decline on the behavioral level. Moreover, results suggests that younger individuals showed higher activation intensity at high CBT load than at low CBT load switching to the opposite pattern at an age of about 40 years. Consistent with the assumptions of the Compensation-Related Utilization of Neural Circuits Hypothesis (CRUNCH), the present results reveal specific age-related alterations in left dorsolateral prefrontal cortex activation in response to increasing task load. Specifically, the results point toward increasing neural inefficiency with age at low task load and a progressive limitation of resources with age at higher task load. The present findings argue for an increasing functional cerebral dysfunction over a time span of 50 years that may partly be compensated on the behavioral level until a resource ceiling is approached.

Trait anxiety and the dynamics of attentional control.

According to recent theoretical approaches dispositional anxiety is fundamentally linked to neural mechanisms of cognitive control (Braver et al., 2007; Eysenck et al., 2007). The present study was conducted to further investigate this topic by focusing on the relation between trait anxiety, conflict-processing and dynamic adjustments in attentional allocation. Participants completed a modified version of the face-word Stroop task while an electroencephalogram was recorded. We analyzed behavioral and electrophysiological correlates of conflict processing and conflict-driven modulations in target and distractor processing. Anxiety was not related to general conflict-sensitivity but to individual differences in conflict-driven adjustments in attentional allocation: following a high level of stimulus-response conflict, highly anxious participants allocated more attentional resources to the processing of predominantly task-relevant information and withdrew attention from the processing of predominantly task-irrelevant information. Thus, trait anxiety appears to be closely related to individual differences in dynamic adjustments of attentional control.

Trait anxiety and dynamic adjustments in conflict processing

Recently, it has been assumed that high- and low-trait-anxious subjects differ in the way they use fundamental cognitive control mechanisms. The present study was designed to further elucidate this topic by focusing on trial-to-trial adjustments in neuronal correlates of conflict processing. An electroencephalogram was recorded while subjects (N=71) performed a gender discrimination version of the Stroop task. The conflict-related N400 of the ERP was influenced by an interaction between trait anxiety and previous trial context: An additional negative-going deflection in the N400 range was observed when the target—distractor pairing of the directly preceding trial was incongruent, but only in highly anxious subjects. Thus, highly anxious subjects appear to more strongly engage neuronal modules involved in conflict monitoring when previously exposed to a high stimulus-response conflict. These results indicate that trait anxiety is crucially linked to the way the cognitive system dynamically adapts to recent demands.

Supramodal neural processing of abstract information conveyed by speech and gesture

Abstractness and modality of interpersonal communication have a considerable impact on comprehension. They are relevant for determining thoughts and constituting internal models of the environment. Whereas concrete object-related information can be represented in mind irrespective of language, abstract concepts require a representation in speech. Consequently, modality-independent processing of abstract information can be expected. Here we investigated the neural correlates of abstractness (abstract vs. concrete) and modality (speech vs. gestures), to identify an abstractness-specific supramodal neural network. During fMRI data acquisition 20 participants were presented with videos of an actor either speaking sentences with an abstract-social [AS] or concrete-object-related content [CS], or performing meaningful abstract-social emblematic [AG] or concrete-object-related tool-use gestures [CG]. Gestures were accompanied by a foreign language to increase the comparability between conditions and to frame the communication context of the gesture videos. Participants performed a content judgment task referring to the person vs. object-relatedness of the utterances. The behavioral data suggest a comparable comprehension of contents communicated by speech or gesture. Furthermore, we found common neural processing for abstract information independent of modality (AS > CS ∩ AG > CG) in a left hemispheric network including the left inferior frontal gyrus (IFG), temporal pole, and medial frontal cortex. Modality specific activations were found in bilateral occipital, parietal, and temporal as well as right inferior frontal brain regions for gesture (G > S) and in left anterior temporal regions and the left angular gyrus for the processing of speech semantics (S > G). These data support the idea that abstract concepts are represented in a supramodal manner. Consequently, gestures referring to abstract concepts are processed in a predominantly left hemispheric language related neural network.

The significance of caudate volume for age-related associative memory decline

Aging comes along with reduced gray matter (GM) volume in several cerebral areas and with cognitive performance decline in different cognitive domains. Moreover, regional GM volume is linked to specific cognitive sub processes in older adults. However, it remains unclear which regional changes in older individuals are directly associated with decreased cognitive performance. Moreover, most of the studies on this topic focused on hippocampal and prefrontal brain regions and their relation to memory and executive functioning. Interestingly, there are only a few studies that reported an association between striatal brain volume and cognitive performance. This is insofar surprising that striatal structures are (1) highly affected by age and (2) involved in different neural circuits that serve intact cognition. To address these issues, voxel-based morphometry (VBM) was used to analyze GM volume in 18 younger and 18 older adults. Moreover, several neuropsychological tests from different neuropsychological test batteries were applied to assess a broad range of cognitive domains. Older adults showed less GM volume than younger adults within frontal, striatal, and cerebellar brain regions. In the group of older adults, significant correlations were found between striatal GM volume and memory performance and between prefrontal/temporal GM volume and executive functioning. The only direct overlap between brain regions associated with regional atrophy and cognitive performance in older adults was found for the right caudate: older adults showed reduced caudate volume relative to younger adults. Moreover, caudate volume was positively correlated with associative memory accuracy in older adults and older adults showed poorer performances than younger adults in the respective associative memory task. Taken together, the current findings indicate the relevance of the caudate for associative memory decline in the aging brain.

Neuroimaging Evidence for Processes Underlying Repetition of Ignored Stimuli

Prolonged response times are observed with targets having been presented as distractors immediately before, called negative priming effect. Among others, inhibitory and retrieval processes have been suggested underlying this behavioral effect. As those processes would involve different neural activation patterns, a functional magnetic resonance imaging (fMRI) study including 28 subjects was conducted. Two tasks were used to investigate stimulus repetition effects. One task focused on target location, the other on target identity. Both tasks are known to elicit the expected response time effects. However, there is less agreement about the relationship of those tasks with the explanatory accounts under consideration. Based on within-subject comparisons we found clear differences between the experimental repetition conditions and the neutral control condition on neural level for both tasks. Hemodynamic fronto-striatal activation patterns occurred for the location-based task favoring the selective inhibition account. Hippocampal activation found for the identity-based task suggests an assignment to the retrieval account; however, this task lacked a behavioral effect.