[no longer on the handout]

Bartels, A., & Zeki, S. (2004). The chronoarchitecture of the human brain - natural viewing conditions reveal a time-based anatomy of the brain. Neuroimage, 22(1), 419-433.

A dominant tendency in cerebral studies has been the attempt to locate architecturally distinct parts of the cortex and assign special functions to each, through histological, clinical or hypothesis-based imaging experiments. Here we show that the cerebral cortex can also be subdivided into different components temporally, without any a priori hypotheses, based on the principle of functional independence. This states that distinct functional subdivisions have activity time courses (ATCs) that are, if not independent, at least characteristic to each when the brain is exposed to natural conditions. To approach a time- based anatomy experimentally, we recorded whole-brain activity using functional magnetic resonance imaging (fMRI) and analyzed the data with independent component analysis (ICA). Our results show that a multitude of cortical areas can be identified based purely on their characteristic ATCs during natural conditions. We demonstrate that a more 'rich' stimulation (free viewing of a movie) leads to more areas being activated in a specific way than conventional stimuli, allowing for a more detailed dissection of the cortex into its subdivisions. We show that stimulus-driven functionally specialized areas can be identified by intersubject correlation even if their function is unknown. Chronoarchitectonic mapping thus opens the prospect of identifying previously unknown cortical subdivisions based on natural viewing conditions by exploiting the characteristic temporal 'fingerprint' that is unique to each. (C) 2004 Elsevier Inc. All rights reserved.

Hennenlotter, A., Schroeder, U., Erhard, P., Haslinger, B., Stahl, R., Weindl, A., von Einsiedel, H. G., Lange, K. W., & Ceballos-Baumann, A. O. (2004). Neural correlates associated with impaired disgust processing in pre-symptomatic Huntington's disease. Brain, 127(6), 1446-1453.

Disturbances in recognizing facial expressions of disgust have been reported previously in pre-symptomatic and manifest Huntington's disease. Given the substantial role of the insula and basal ganglia in the perception of disgust as revealed by functional imaging, lesion studies and intracerebral recordings, we propose dysfunction within the insula and/or basal ganglia as the underlying neural substrate. Using functional MRI (fMRI), we studied a group of nine pre-symptomatic Huntington's disease gene carriers and nine healthy controls, matched for age, gender, intelligence and years of education, while they were viewing disgusted facial expressions. As control conditions, surprised and neutral expressions were presented. Compared with healthy controls, Huntington's disease gene carriers showed reduced responses within the left dorsal anterior insula during processing of disgusted facial expressions. Moreover, processing of disgust was associated with significant activation of the left dorsal anterior insula and putamen in healthy controls, but not in Huntington's disease gene carriers. Furthermore, behavioural assessment revealed a selective impairment in recognizing facial expressions displaying disgust in Huntington's disease gene carriers. Our finding of dysfunctional decreased insula activation in pre-symptomatic Huntington's disease provides an explanation for the clinical deficit in recognizing facial expression of disgust. Furthermore, it underscores the role of the insula in the emotion of disgust.

Liegeois, F., Connelly, A., Cross, J. H., Boyd, S. G., Gadian, D. G., Vargha-Khadem, F., & Baldeweg, T. (2004). Language reorganization in children with early-onset lesions of the left hemisphere: an fMRI study. Brain, 127(6), 1229-1236.

It is widely assumed that following extensive damage to the left hemisphere sustained in early childhood, language functions are likely to reorganize and develop in the right hemisphere, especially if the lesion affects the classical Broca's or Wernicke's language areas. In the present study, functional MRI (fMRI) was used to examine language lateralization in 10 children and adolescents with intractable epilepsy who sustained an early lesion in the left hemisphere. Lesions were adjacent to or within anterior language cortex in five patients, while they were remote from both Broca's and Wernicke's areas in the remainder. A lateralization index was calculated on the basis of the number of voxels activated in the left and right inferior frontal gyri when performing a covert verb generation task. Only two patients were right-handed, suggesting a high incidence of functional reorganization for motor control in the remaining patients. Five out of 10 showed bilateral or right language lateralization, but lateralization could not be inferred from the proximity of lesions to classical language areas on an individual basis. Lesions in or near Broca's area were not associated with inter-hemispheric language reorganization in four out of five cases, but with perilesional activation within the damaged left hemisphere. Paradoxically, lesions remote from the classical language areas were associated with non-left language lateralization in four out of five cases. Finally, handedness, age at onset of chronic seizures, and site of EEG abnormality also showed no obvious association with language lateralization. In conclusion, it is difficult to infer intra- versus inter-hemispheric language reorganization on the basis of clinical observations in the presence of early pathology to the left hemisphere.

Matsumoto, R., Nair, D. R., LaPresto, E., Najm, I., Bingaman, W., Shibasaki, H., & Luders, H. O. (2004). Functional connectivity in the human language system: a cortico-cortical evoked potential study. Brain, 127(10), 2316-2330.

A better understanding of the mechanisms involved in human higher cortical functions requires a detailed knowledge of neuronal connectivity between functional cortical regions. Currently no good method for tracking in vivo neuronal connectivity exists. We investigated the inter-areal connections in vivo in the human language system using a new method, which we termed cortico-cortical evoked potentials' (CCEPs). Eight patients with epilepsy (age 13-42 years) underwent invasive monitoring with subdural electrodes for epilepsy surgery. Six patients had language dominance on the side of grid implantation and two had bilateral language representation by the intracarotid amobarbital test. Conventional cortical electrical stimulation was performed to identify the anterior and posterior language areas. Single pulse electrical stimuli were delivered to the anterior language (eight patients), posterior language (four patients) or face motor (two patients) area, and CCEPs were obtained by averaging electrocorticograms (ECoGs) recorded from the perisylvian and extrasylvian basal temporal language areas time-locked to the stimulus. The subjects were not asked to perform any tasks during the study. Stimulation at the anterior language area elicited CCEPs in the lateral temporo-parietal area (seven of eight patients) in the middle and posterior part of the superior temporal gyrus, the adjacent part of the middle temporal gyrus and the supramarginal gyrus. CCEPs were recorded in 3-21 electrodes per patient. CCEPs occurred at or around the particular electrodes in the posterior language area which, when stimulated, produced speech arrest. Similar early and late CCEPs were obtained from the basal temporal area by stimulating the anterior language area (three of three patients). In contrast, stimulation of the adjacent face motor area did not elicit CCEPs in language areas but rather in the postcentral gyrus. Stimulation of the posterior language area produced CCEPs in the anterior language (three of four patients) as well as in the basal temporal area (one of two patients). These CCEPs were less well defined. These findings suggest that perisylvian and extrasylvian language areas participate in the language system as components of a network by means of feed-forward and feed-back projections. Different from the classical Wernicke-Geschwind model, the present study revealed a bidirectional connection between Broca's and Wernicke's areas probably through the arcuate fasciculus and/or the cortico-subcortico-cortical pathway. CCEPs were recorded from a larger area than the posterior language area identified by electrical stimulation. This suggests the existence of a rather broad neuronal network surrounding the previously recognized core region of this area.

Ruby, P., & Decety, J. (2004). How would you feel versus how do you think she would feel? A neuroimaging study of perspective-taking with social emotions. Journal of Cognitive Neuroscience, 16(6), 988-999.

Perspective-taking is a complex cognitive process involved in social cognition. This positron emission tomography (PET) study investigated by means of a factorial design the interaction between the emotional and the perspective factors. Participants were asked to adopt either their own (first person) perspective or the (third person) perspective of their mothers in response to situations involving social emotions or to neutral situations. The main effect of third-person versus first- person perspective resulted in hemodynamic increase in the medial part of the superior frontal gyrus, the left superior temporal sulcus, the left temporal pole, the posterior cingulate gyrus, and the right inferior parietal lobe, A cluster in the postcentral gyrus was detected in the reverse comparison. The amygdala was selectively activated when subjects were processing social emotions, both related to self and other. Interaction effects were identified in the left temporal pole and in the right postcentral gyrus. These results support our prediction that the frontopolar, the somatosensory cortex, and the right inferior parietal lobe are crucial in the process of self/ other distinction. In addition, this study provides important building blocks in our understanding of social emotion processing and human empathy.

Schendel, K., & Robertson, L. C. (2004). Reaching out to see: Arm position can attenuate human visual loss. Journal of Cognitive Neuroscience, 16(6), 935-943.

Electrophysiological recordings in monkeys have now revealed several brain regions that contain bimodal visuotactile neurons capable of responding to either tactile or visual stimuli placed on or near the hands, arms, and face. These cells have now been found in frontal, parietal, and subcortical areas of the monkey brain, suggesting a cortical network of neurons that preferentially represent near peripersonal space. The degree to which the visual responses of such cells rely on input from the primary visual cortex and the extent to which they may contribute to visual perception is not completely understood. Nonetheless, recent neuropsychological studies suggest that a similar representation of near space may be bimodally coded in humans as well, Given the accumulating evidence for specialized processing of visual stimuli placed near the hands and arms, we hypothesized that arm position may be capable of modulating human visual ability. Here we report the case of WM, who lost his ability to see in his left visual hemifield after sustaining damage to his right primary visual cortex. Interestingly, the placement of WM's left arm into his "blind" field resulted in significantly better detection of left visual field stimuli compared to when his hand was placed in his lap at midline. Moreover, we found this attenuation to be confined to stimuli presented within reaching distance (unless a tool that extended WM's reach was held while lie performed the test). These findings are highly consistent with the characteristics of the bimodal visuo-tactile neurons that have been described in monkeys, Thus, it seems that arm position can modulate human visual ability, even after damage to the primary visual cortex. This study provides an exciting bridge between monkey neurophysiology and human visual capacity While also offering a novel approach for improving visual defects acquired via cortical injury.