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ABSTRACT: Planning of behavior relies on the integrity of the mid-dorsolateral prefrontal cortex (mid-dlPFC). Yet, only indirect evidence exists on the association of protracted maturation of dlPFC and continuing gains in planning performance post adolescence. Here, gray matter density of mid-dlPFC in young, healthy adults (18-32years) was regressed onto performance on the Tower of London planning task while accounting for moderating effects of age and sex on this interrelation. Multiple regression analysis revealed an association of planning performance and mid-dlPFC gray matter density that was especially strong in late adolescence and early twenties. As expected, for males better planning performance was linked to reduced gray matter density of mid-dlPFC, possibly due to maturational processes such as synaptic pruning. Most surprisingly, females showed an inverted, positive interrelation of planning performance and mid-dlPFC gray matter density, indicating that sexually dimorphic development of dlPFC continues during early adulthood. Age and sex are hence important moderators of the link between planning performance and gray matter density in mid-dlPFC. Consequently, the assessment of moderator effects in regression designs can significantly enhance understanding of brain-behavior relationships.
NeuroImage 08/2012; 63(3):1454-63. · 5.89 Impact Factor
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ABSTRACT: We used an identical repetition priming paradigm in functional MRI (fMRI) and magnetoencephalography (MEG) to investigate brain networks modulated by stimulus repetition and familiarity. In particular, pictures of familiar or unfamiliar objects were presented sequentially, with stimulus repetitions occurring within few trials. The results of both studies indicated close agreement between the pattern found in fMRI-BOLD (blood oxygenation level dependent) responses and in source localizations of induced gamma-band activity derived from MEG. In both studies, the brain regions that were significantly associated with repetition suppression in response to familiar visual objects encompassed bilaterally the medial and lateral occipital cortex, inferior occipitotemporal regions including the left fusiform cortex, as well as parietal areas. Modulations by stimulus familiarity occurred mainly within this network. Overall, we found noticeable correspondences between the results of fMRI-BOLD signals and MEG gamma-band activity, suggesting that both methods can be used in analogous ways to study the neural basis of repetition priming and object recognition.
Neuroreport 07/2012; 23(13):757-61. · 1.66 Impact Factor
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ABSTRACT: In a companion study, eye-movement analyses in the Tower of London task (TOL) revealed independent indicators of functionally separable cognitive processes during problem solving, with processes of building up an internal representation of the problem preceding actual planning processes. These results imply that processes of internalization and planning should also be distinguishable in time and space with respect to concomitant brain activation patterns. To investigate this possibility, here we conducted analyses of fMRI data for left and right dorsolateral prefrontal cortex (dlPFC) during problem solving in the TOL task by accounting for the trial-by-trial variability of onsets and durations of the different cognitive processing stages. Comparisons between stimulus-locked and response-locked modeling approaches affirmed that activation in left dlPFC was elicited particularly during early processes of internalization, comprising the extraction of goal information and the generation of an internal problem representation, whereas activation in right dlPFC was predominantly attributable to later processes of mental transformations on this representation, that is planning proper. Thus, present data corroborate the proposal that often observed bilateral dlPFC activation patterns during complex cognitive tasks such as problem solving may reflect functionally and, to some extent, even temporally separable processes with opposing lateralizations.
Brain and Cognition 03/2012; 80(1):170-6. · 3.17 Impact Factor
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ABSTRACT: The role of verbal and visuospatial information processing in Tower of London (TOL) tasks was investigated. The first part of the investigation examined the verbal and visuospatial abilities and preferred cognitive style (visualizer vs. verbalizer) of 79 participants, in an inter-individual differences approach. Visuospatial abilities significantly predicted TOL performance, but the impact of cognitive style was negligible. The second part applied a dual-task manipulation of concurrent interference of TOL planning tasks on verbal and visuospatial memory, using the same participants. Concurrent processing of the TOL tasks diminished visuospatial memory performance considerably but had no effect on verbal memory, and there was no interaction between cognitive style and memory. These findings clearly underscore the role of visuospatial information processing in TOL tasks and indicate little bearing of verbal or visual cognitive style on TOL problem solving. These results have important implications for TOL and cognitive style in clinical application and cognitive neuroimaging research.
British Journal of Psychology 02/2012; 103(1):98-116. · 2.37 Impact Factor
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ABSTRACT: Most neuroimaging studies on planning report bilateral activations of the dorsolateral prefrontal cortex (dlPFC). Recently, these concurrent activations of left and right dlPFC have been shown to double dissociate with different cognitive demands imposed by the planning task: Higher demands on the extraction of task-relevant information led to stronger activation in left dlPFC, whereas higher demands on the integration of interdependent information into a coherent action sequence entailed stronger activation of right dlPFC. Here, we used continuous theta-burst stimulation (cTBS) to investigate the supposed causal structure-function mapping underlying this double dissociation. Two groups of healthy subjects (left-lateralized stimulation, n = 26; right-lateralized stimulation, n = 26) were tested within-subject on a variant of the Tower of London task following either real cTBS over dlPFC or sham stimulation over posterior parietal cortex. Results revealed that, irrespective of specific task demands, cTBS over left and right dlPFC was associated with a global decrease and increase, respectively, in initial planning times compared to sham stimulation. Moreover, no interaction between task demands and stimulation type (real vs. sham) and/or stimulation side (left vs. right hemisphere) were found. Together, against expectations from previous neuroimaging data, lateralized cTBS did not lead to planning-parameter specific changes in performance, but instead revealed a global asymmetric pattern of faster versus slower task processing after left versus right cTBS. This global asymmetry in the absence of any task-parameter specific impact of cTBS suggests that different levels of information processing may span colocalized, but independent axes of functional lateralization in the dlPFC. Hum Brain Mapp, 2011. © 2011 Wiley Periodicals, Inc.
Human Brain Mapping 10/2011; · 5.88 Impact Factor
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ABSTRACT: Working memory supports the recognition of objects in the environment. Memory models have postulated that recognition relies on 2 processes: assessing the degree of similarity between an external stimulus and memory representations and testing the resulting summed-similarity value against a critical level for recognition. Here, we varied the similarity between samples held in working memory and a probe to investigate these 2 processes with magnetoencephalography. Two separable components matched our expectations: First, from 280 ms after probe onset, clearly nonmatching probes differed from both similar nonmatches and matches over left frontal cortex. At 350-400 ms, these signals evolved into a pattern of gradually increasing activation as a function of sample-probe similarity, as expected for a neural representation of summed similarity. Second, a signal potentially reflecting criterion testing was observed at 600-700 ms at right frontotemporal sensors that differentiated between matches and nonmatches without further differences between similar and dissimilar probes. Thus, analysis of the time course of recognition provided strong evidence that similarity summation and criterion testing have separable neural bases. As probably both working and long-term memory recognition draw on these processes, they may be involved in many domains of behavior.
Cerebral Cortex 09/2011; 22(8):1950-8. · 6.54 Impact Factor
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ABSTRACT: The analysis of cortical and subcortical networks requires the identification of their nodes, and of the topology and dynamics of their interactions. Exploratory tools for the identification of nodes are available, e.g. magnetoencephalography (MEG) in combination with beamformer source analysis. Competing network topologies and interaction models can be investigated using dynamic causal modelling. However, we lack a method for the exploratory investigation of network topologies to choose from the very large number of possible network graphs. Ideally, this method should not require a pre-specified model of the interaction. Transfer entropy--an information theoretic implementation of Wiener-type causality--is a method for the investigation of causal interactions (or information flow) that is independent of a pre-specified interaction model. We analysed MEG data from an auditory short-term memory experiment to assess whether the reconfiguration of networks implied in this task can be detected using transfer entropy. Transfer entropy analysis of MEG source-level signals detected changes in the network between the different task types. These changes prominently involved the left temporal pole and cerebellum--structures that have previously been implied in auditory short-term or working memory. Thus, the analysis of information flow with transfer entropy at the source-level may be used to derive hypotheses for further model-based testing.
Progress in Biophysics and Molecular Biology 03/2011; 105(1-2):80-97. · 3.20 Impact Factor
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ABSTRACT: It is well established that the mid-dorsolateral prefrontal cortex (dlPFC) plays a critical role in planning. Neuroimaging studies have yielded predominantly bilateral dlPFC activations, but the existence and nature of functionally specific contributions of left and right dlPFC have remained elusive. In recent experiments, 2 independent parameters have been identified which substantially determine planning: 1) the degree of interdependence between consecutive steps (search depth) and 2) the degree to which the configuration of the goal state renders the order of single steps either clearly evident or ambiguous (goal hierarchy). Thus, search depth affects the actual mental generation and evaluation of action sequences, whereas goal hierarchy reflects the extraction of goal information from an encountered problem. Here, both parameters were independently manipulated in an event-related functional magnetic resonance imaging study using the Tower of London task. Results revealed a double dissociation as indicated by a significant crossover interaction of hemisphere and task parameter: in left dlPFC, activations were stronger for higher demands on goal hierarchy than on search depth, whereas the reversed result emerged in right dlPFC. In conclusion, often observed bilateral patterns of dlPFC activation in complex tasks may reflect the concomitant operation of specific cognitive processes that show opposing lateralizations.
Cerebral Cortex 02/2011; 21(2):307-17. · 6.54 Impact Factor
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ABSTRACT: In a previous study (Unterrainer, Kaller, Halsband, & Rahm, 2006), chess players outperformed non-chess players in the Tower of London planning task but exhibited disproportionately longer processing times. This pattern of results raises the question of whether chess players' planning capabilities are superior or whether the results reflect differences in the speed-accuracy trade-off between the groups, possibly attributable to sports motivation. The present study was designed to disambiguate these alternative suggestions by implementing various constraints on planning time and by assessing self-reported motivation. In contrast to the previous study, chess players' performance was not superior, independently of whether problems had to be solved with (Experiment 1) or without (Experiment 2) time limits. As expected, chess players reported higher overall trait and state motivation scores across both experiments. These findings revise the notion of superior planning performance in chess players. In consequence, they do not conform with the assumption of a general transfer of chess-related planning expertise to other cognitive domains, instead suggesting that superior performance may be possible only under specific circumstances such as receiving competitive instructions.
The American Journal of Psychology 01/2011; 124(2):213-25. · 1.09 Impact Factor
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ABSTRACT: Human gamma-band activity (GBA) has been related to a variety of functions ranging from perception and attention to memory and consciousness. Indeed fast spectral activity derived during numerous experimental paradigms has been interpreted as providing support for the functional importance of these signals. The present review provides an overview of findings demonstrating direct or indirect associations between GBA and behavioral measures of task performance or perceptual experience in humans. While the majority of papers have focused on perception and awareness, relationships between GBA and behavior have also been observed during insightful problem solving, short- or long-term memory, and motor tasks. In these studies, GBA was reported to predict behavioral measures such as correct response rates or reaction times both in healthy subjects and patients with neuropsychiatric disorders. The review demonstrates that there is increasingly strong evidence for a close association between GBA during both perceptual and cognitive tasks and behavioral outcome measures. With this in mind, the investigation of GBA might help elucidate the mechanisms underlying deficient functioning in patients with neuropsychiatric disorders.
International journal of psychophysiology: official journal of the International Organization of Psychophysiology 01/2011; 79(1):39-48. · 3.05 Impact Factor
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ABSTRACT: Cognitive, clinical, and neuroimaging studies on planning abilities most frequently implement the Tower of London task or one of its variants. Yet, cumulating evidence from a series of experiments suggests that the commonly used approximation of problem difficulty in terms of the minimum number of moves for goal attainment is too coarse a measure for the underlying cognitive operations, and in some cases may be even misleading. Rather, problem difficulty can be more specifically characterized by a set of structural task parameters such as the number and nature of optimal and suboptimal solution paths, the required search depths, the patterns of intermediate and goal moves, goal hierarchies and the associated degree of ambiguity in the sequential ordering of goal moves. First applications in developmental and patient studies have proven fruitful in targeting fundamental alterations of planning abilities in healthy and clinical conditions. In addition, recent evidence from neuroimaging shows that manipulations of problem structure relate to separate cognitive and neural processes and are accompanied by dissociable brain activation patterns. Here, we briefly review these structural problem parameters and the concepts behind. As controlling for task parameters and selecting a balanced problem set is a complex and error-prone endeavor, we further present TowerTool, a software solution that allows easy access to in-depth analysis of the problem structure of widely used planning tasks like the Tower of London, the Tower of Hanoi, and their variants. Thereby, we hope to encourage and facilitate the implementation of structurally balanced task sets in future studies on planning and to promote transfer between the cognitive, developmental, and clinical neurosciences.
Behavioural brain research 01/2011; 216(1):1-8. · 3.22 Impact Factor
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ABSTRACT: Working memory (WM) constitutes a fundamental aspect of human cognition. It refers to the ability to keep information active for further use, while allowing it to be prioritized, modified and protected from interference. Much research has addressed the storage function of WM, however, its 'working' aspect still remains underspecified. Many operations that work on the contents of WM do not appear specific to WM. The present review focuses on those operations that we consider "basic" because they operate in the service of memory itself, by providing its basic functionality of retaining information active, in a stable yet flexible way. Based on current process models of WM we review five strands of research: (1) mnemonic selection of one item amongst others, (2) updating the focus of attention with the selected item, (3) updating the content of visual WM with new item(s), (4) rehearsal of visuospatial information and (5) coping with interference. We discuss the neuronal substrates underlying those operations obtained with functional magnetic resonance imaging and relate them to findings on "executive functions". The presented data support the view that WM emerges from interactions between higher sensory, attentional and mnemonic functions, with separable neural bases. However, interference processing and the representation of rule switching in WM may demand an extension of the current WM models by executive control functions.
Behavioural brain research 12/2010; 214(2):172-9. · 3.22 Impact Factor
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ABSTRACT: Decoding of frequency-modulated (FM) sounds is essential for phoneme identification. This study investigates selectivity to FM direction in the human auditory system.
Magnetoencephalography was recorded in 10 adults during a two-tone adaptation paradigm with a 200-ms interstimulus-interval. Stimuli were pairs of either same or different frequency modulation direction. To control that FM repetition effects cannot be accounted for by their on- and offset properties, we additionally assessed responses to pairs of unmodulated tones with either same or different frequency composition. For the FM sweeps, N1m event-related magnetic field components were found at 103 and 130 ms after onset of the first (S1) and second stimulus (S2), respectively. This was followed by a sustained component starting at about 200 ms after S2. The sustained response was significantly stronger for stimulation with the same compared to different FM direction. This effect was not observed for the non-modulated control stimuli.
Low-level processing of FM sounds was characterized by repetition enhancement to stimulus pairs with same versus different FM directions. This effect was FM-specific; it did not occur for unmodulated tones. The present findings may reflect specific interactions between frequency separation and temporal distance in the processing of consecutive FM sweeps.
PLoS ONE 01/2010; 5(12):e15548. · 4.09 Impact Factor
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ABSTRACT: Cognition depends critically on working memory, the active representation of a limited number of items over short periods of time. In addition to the maintenance of information during the course of cognitive processing, many tasks require that some of the items in working memory become transiently more important than others. Based on cognitive models of working memory, we hypothesized two complementary essential cognitive operations to achieve this: a selection operation that retrieves the most relevant item, and an updating operation that changes the focus of attention onto it. Using functional magnetic resonance imaging, high-resolution oculometry, and behavioral analysis, we demonstrate that these two operations are functionally and neuroanatomically dissociated. Updating the attentional focus elicited transient activation in the caudal superior frontal sulcus and posterior parietal cortex. In contrast, increasing demands on selection selectively modulated activation in rostral superior frontal sulcus and posterior cingulate/precuneus. We conclude that prioritizing one memory item over others invokes independent mechanisms of mnemonic retrieval and attentional focusing, each with its distinct neuroanatomical basis within frontal and parietal regions. These support the developing understanding of working memory as emerging from the interaction between memory and attentional systems.
Journal of Neuroscience 10/2009; 29(43):13735-41. · 7.11 Impact Factor
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ABSTRACT: Identifying overtly observable indicators of cognitive processes should provide a promising basis for a more precise tracking of the associated cognitive and neural events. In the current study we used recordings of eye movements to gain deeper insight into the time course of visuospatial problem solving as measured by the Tower of London. Single-trial, saccade-locked analyses revealed that, despite the complexity of the implemented task, gaze alternations between start and goal state followed a highly regular pattern. Consistent with the buildup of an internal representation, the first two fixations were of constant duration and unaffected by experimental variations of planning demands. Instead, planning manipulations exclusively influenced the duration of the very last fixation before problem execution. Our results demonstrate that different phases of complex cognition can be identified on a single-trial level using eye movement analyses.
Psychophysiology 05/2009; 46(4):818-30. · 3.29 Impact Factor
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ABSTRACT: The short-term retention of information has been related to oscillatory activity in the gamma-band. In recent auditory spatial short-term memory studies we have found stimulus-specific components of parieto-occipital gamma-band activity (GBA) which might reflect the activation of local networks tuned to task-relevant stimulus features. The present magnetoencephalography study (N=22) tested this interpretation by assessing whether the topography of stimulus-specific GBA depends on task demands. Sample sounds were characterized by both a variable interaural time delay and a variable central frequency. In separate task blocks, either the lateralization or the frequency of the same stimuli had to be maintained. Statistical probability mapping of differences in oscillatory responses to the retention of sample sounds replicated the contralateral posterior topography for GBA components distinguishing between medial and lateral sounds in the spatial memory task. In contrast, lower- and higher-frequency stimuli were accompanied by frontal GBA components in the frequency task. Memory for lateralization versus frequency selectively enhanced oscillatory activity for these posterior versus frontal components, directly demonstrating their modulation by task demands. Incorrect "non-match" responses were negatively correlated with delay-phase GBA to the relevant feature, whereas incorrect "match" responses correlated positively with GBA to the irrelevant feature. In summary, the topography of stimulus-specific GBA to identical stimuli reflected the selective representation of task-relevant features. Task performance was predicted by both enhanced stimulus-specific GBA for the task-relevant stimulus attribute and reduced gamma-band representations of the task-irrelevant stimulus feature. Stimulus-specific GBA may reflect the memory representation that is used in subsequent recognition.
NeuroImage 04/2009; 46(4):1127-36. · 5.89 Impact Factor
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ABSTRACT: Recently we have demonstrated that during auditory short-term memory maintenance, gamma-band activity (GBA) components can be identified which are specific to the retained stimulus. These activations peaked in the middle of the delay phase between sample and test stimuli, and their magnitude during the final part of this period correlated with performance. However, using a constant delay duration did not allow to answer the question whether stimulus-specific GBA components represented responses to sample sounds or anticipatory activations preceding test stimuli. Here we addressed this unresolved issue by investigating the temporal dynamics of stimulus-specific GBA during two delay durations. Magnetoencephalogram was recorded in 18 adults during an auditory spatial short-term memory task involving lateralized sample stimuli presented with two different interaural time delays. Subjects had to decide whether test stimuli presented after retention phases of 800 or 1200 ms had the same lateralization as sample sounds. Statistical probability mapping served to identify oscillatory activations differentiating between the two sample sounds. We found stimulus-specific GBA components over posterior cortex peaking about 400 ms prior to the onset of test stimuli regardless of delay duration. Their magnitude correlated with task performance. In summary, stimulus-specific GBA components with a predictive power for short-term memory performance were observed in anticipation of test stimuli. They may reflect the preparatory activation of memory representations or the shifting of attention to the specific expected location of the test stimulus.
NeuroImage 09/2008; 44(1):257-64. · 5.89 Impact Factor
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ABSTRACT: The ability to plan and search ahead is essential for problem solving in most situations in everyday life. To investigate the development of planning and related processes, a sample of four- and five-year-old children was examined in a variant of the Tower of London, a frequently used neuropsychological assessment tool of planning abilities. The applied problems either required searching ahead for optimal solution or were solvable by pure step-by-step forward processing. Furthermore, the ambiguity of subgoal ordering was varied. Results revealed an age-related effect of search depth: the four-year olds' planning accuracy was particularly decreased in problems demanding search ahead, while five-year olds mastered both problem types equally well. Interestingly, this interaction between age and search depth could not be accounted for by measures of working memory and inhibition. Differential effects of age were also found for subgoal ordering with respect to initial planning and movement execution times. In sum, planning abilities showed considerable development during late kindergarten age that appeared to be specifically associated with the integration and back-validation of the anticipated consequences of internally modeled actions. The present study demonstrates that a careful consideration of problem structure may greatly enhance the insights gained from the application of a routinely used assessment tool, the Tower of London. This may be especially advantageous when addressing specific subpopulations such as children or clinical samples.
Brain and Cognition 09/2008; 67(3):360-70. · 3.17 Impact Factor
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ABSTRACT: Contrasting electro- or magnetoencephalographic oscillatory responses to sensory stimuli with a pre-stimulation baseline commonly yields spectrally broad and topographically distributed activations. In contrast, comparisons between closely matched task conditions usually result in more focal differences. In the present study, we reanalyzed an existing set of MEG data recorded during stimulation with virtual Kanizsa figures and no-triangle control stimuli to contrast results yielded by the two approaches. Statistical analysis showed that visual stimulation compared to baseline gave rise to spectral amplitude reductions in lower frequencies including alpha and beta and amplitude enhancements in gamma frequencies above 55 Hz. These changes reached significance by about 100 ms post-stimulus onset, were topographically widespread over posterior cortex, and did not differ between stimuli. A second, more focal component over ventral occipital cortex peaked at about 300 ms in the gamma range at approximately 70 Hz. It was more pronounced for the Kanizsa triangle than for the no-triangle stimulus. A third gamma component over lateral occipito-temporal cortex showed an amplitude increase at around 450 ms for virtual figures and a concomitant decrease for the nongestalt-like control stimulus, and no overall task-related activity. Our findings illustrate that direct comparisons between conditions yield effects with a more focal spectral and topographical distribution than comparisons with a pre-stimulus baseline. Moreover, they exemplify that contrasts between conditions may reveal additional activations not captured by comparisons with a pre-stimulus baseline.
Brain Research 07/2008; 1235:63-73. · 2.73 Impact Factor
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ABSTRACT: The human amygdala is thought to play a pivotal role in the processing of emotionally significant sensory information. The major subdivisions of the human amygdala-the laterobasal group (LB), the superficial group (SF), and the centromedial group (CM)-have been anatomically delineated, but the functional response properties of these amygdala subregions in humans are still unclear. We combined functional MRI with cyto-architectonically defined probabilistic maps to analyze the response characteristics of amygdala subregions in subjects presented with auditory stimuli. We found positive auditory stimulation-related signal changes predominantly in probabilistically defined LB, and negative responses predominantly in SF and CM. In the left amygdala, mean response magnitude in the core area of LB with 90-100% assignment probability was significantly larger than in the core areas of SF and CM. These differences were observed for pleasant and unpleasant stimuli. Our findings reveal that the probabilistically defined anatomical subregions of the human amygdala show distinctive fMRI response patterns. The stronger auditory responses in LB as compared with SF and CM may reflect a predominance of auditory inputs to human LB, similar to many animal species in which the majority of sensory, including auditory, afferents project to this subdivision of the amygdala. Our study indicates that the intrinsic functional differentiation of the human amygdala may be probed using fMRI combined with probabilistic anatomical maps.
PLoS ONE 02/2007; 2(3):e307. · 4.09 Impact Factor
