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1.  Altered network properties of the fronto-parietal network and the thalamus in impaired consciousness☆ 
NeuroImage : Clinical  2013;4:240-248.
Recovery of consciousness has been associated with connectivity in the frontal cortex and parietal regions modulated by the thalamus. To examine this model and to relate alterations to deficits in cognitive functioning and conscious processing, we investigated topological network properties in patients with chronic disorders of consciousness recovered from coma.
Resting state fMRI data of 34 patients with unresponsive wakefulness syndrome and 25 in minimally conscious state were compared to 28 healthy controls. We investigated global and local network characteristics. Additionally, behavioral measures were correlated with the local metrics of 28 regions within the fronto-parietal network and the thalamus.
In chronic disorders of consciousness, modularity at the global level was reduced suggesting a disturbance in the optimal balance between segregation and integration. Moreover, network properties were altered in several regions which are associated with conscious processing (particularly, in medial parietal, and frontal regions, as well as in the thalamus). Between minimally conscious and unconscious patients the local efficiency of medial parietal regions differed. Alterations in the thalamus were particularly evident in non-conscious patients. Most of the regions affected in patients with impaired consciousness belong to the so-called ‘rich club’ of highly interconnected central nodes. Disturbances in their topological characteristics have severe impact on information integration and are reflected in deficits in cognitive functioning probably leading to a total breakdown of consciousness.
Highlights
•We investigated network properties in patients with a disorder of consciousness.•Patients showed reduced global modularity.•Alterations in regions of the rich club were related to impaired consciousness.•These alterations have severe impact on information integration and segregation.•Disturbances in overall integration may lead to breakdown of consciousness.
doi:10.1016/j.nicl.2013.12.005
PMCID: PMC3895618  PMID: 24455474
DOC, disorders of consciousness; ACC, anterior cingulate cortex; PCC, posterior cingulate cortex; MCS, minimally conscious state; VS/UWS, vegetative state/unresponsive wakefulness syndrome; Consciousness; Vegetative state; Network; Graph theory; Connectivity; Small world
2.  Processing counterfactual and hypothetical conditionals: An fMRI investigation 
Neuroimage  2013;72(C):265-271.
Counterfactual thinking is ubiquitous in everyday life and an important aspect of cognition and emotion. Although counterfactual thought has been argued to differ from processing factual or hypothetical information, imaging data which elucidate these differences on a neural level are still scarce. We investigated the neural correlates of processing counterfactual sentences under visual and aural presentation. We compared conditionals in subjunctive mood which explicitly contradicted previously presented facts (i.e. counterfactuals) to conditionals framed in indicative mood which did not contradict factual world knowledge and thus conveyed a hypothetical supposition. Our results show activation in right occipital cortex (cuneus) and right basal ganglia (caudate nucleus) during counterfactual sentence processing. Importantly the occipital activation is not only present under visual presentation but also with purely auditory stimulus presentation, precluding a visual processing artifact. Thus our results can be interpreted as reflecting the fact that counterfactual conditionals pragmatically imply the relevance of keeping in mind both factual and supposed information whereas the hypothetical conditionals imply that real world information is irrelevant for processing the conditional and can be omitted. The need to sustain representations of factual and suppositional events during counterfactual sentence processing requires increased mental imagery and integration efforts. Our findings are compatible with predictions based on mental model theory.
Highlights
► Neural correlates of processing counterfactual conditionals were investigated. ► We presented sentences in auditory and visual modality. ► Counterfactuals show stronger activations in cuneus and caudate nucleus. ► The findings are consistent with predictions based on mental model theory.
doi:10.1016/j.neuroimage.2013.01.060
PMCID: PMC3610017  PMID: 23380169
Counterfactual thinking; Conditionals; Subjunctive mood; Indicative mood; fMRI
3.  Opposite effects of visual and auditory word-likeness on activity in the visual word form area 
The present fMRI study investigated the effects of word-likeness of visual and auditory stimuli on activity along the ventral visual stream. In the context of a one-back task, we presented visual and auditory words, pseudowords, and artificial stimuli (i.e., false-fonts and reversed-speech, respectively). Main findings were regionally specific effects of word-likeness on activation in a left ventral occipitotemporal region corresponding to the classic localization of the Visual Word Form Area (VWFA). Specifically, we found an inverse word-likeness effect for the visual stimuli in the form of decreased activation for words compared to pseudowords which, in turn, elicited decreased activation compared to the artificial stimuli. For the auditory stimuli, we found positive word-likeness effects as both words and pseudowords elicited more activation than the artificial stimuli. This resulted from a marked deactivation in response to the artificial stimuli and no such deactivation for words and pseudowords. We suggest that the opposite effects of visual and auditory word-likeness on VWFA activation can be explained by assuming the involvement of visual orthographic memory representations. For the visual stimuli, these representations reduce the coding effort as a function of word-likeness. This results in highest activation to the artificial stimuli and least activation to words for which corresponding representations exist. The positive auditory word-likeness effects may result from activation of orthographic information associated with the auditory words and pseudowords. The view that the VWFA has a primarily visual function is supported by our findings of high activation to the visual artificial stimuli (which have no phonological or semantic associations) and deactivation to the auditory artificial stimuli. According to the phenomenon of cross-modal sensory suppression such deactivations during demanding auditory processing are expected in visual regions.
doi:10.3389/fnhum.2013.00491
PMCID: PMC3756304  PMID: 24009569
fMRI; neuroimaging; one-back task; word-likeness; word processing; VWFA; orthographic representations
4.  Common brain areas engaged in false belief reasoning and visual perspective taking: a meta-analysis of functional brain imaging studies 
We performed a quantitative meta-analysis of functional neuroimaging studies to identify brain areas which are commonly engaged in social and visuo-spatial perspective taking. Specifically, we compared brain activation for visual-perspective taking to activation for false belief reasoning, which requires awareness of perspective to understand someone's mistaken belief about the world which contrasts with reality. In support of a previous account by Perner and Leekam (2008), our meta-analytic conjunction analysis found common activation for false belief reasoning and visual perspective taking in the left but not the right dorsal temporo-parietal junction (TPJ). This fits with the idea that the left dorsal TPJ is responsible for representing different perspectives in a domain-general fashion. Moreover, our conjunction analysis found activation in the precuneus and the left middle occipital gyrus close to the putative Extrastriate Body Area (EBA). The precuneus is linked to mental-imagery which may aid in the construction of a different perspective. The EBA may be engaged due to imagined body-transformations when another's viewpoint is adopted.
doi:10.3389/fnhum.2013.00712
PMCID: PMC3814428  PMID: 24198773
neuroimaging meta-analysis; theory of mind; false belief; visual perspective taking; temporo-parietal junction
5.  A Dual-Route Perspective on Poor Reading in a Regular Orthography: An fMRI Study 
This study examined functional brain abnormalities in dyslexic German readers who – due to the regularity of German in the reading direction – do not exhibit the reading accuracy problem of English dyslexic readers, but suffer primarily from a reading speed problem. The in-scanner task required phonological lexical decisions (i.e., Does xxx sound like an existing word?) and presented familiar and unfamiliar letter strings of existing phonological words (e.g., Taxi-Taksi) together with nonwords (e.g., Tazi). Dyslexic readers exhibited the same response latency pattern (words < pseudohomophones < nonwords) as nonimpaired readers, but latencies to all item types were much prolonged. The imaging results were suggestive for a different neural organization of reading processes in dyslexic readers. Specifically, dyslexic readers, in response to lexical route processes, exhibited underactivation in a left ventral occipitotemporal region which presumably is engaged by visual-orthographic whole word recognition. This region was also insensitive to the increased visual-orthographic processing demands of the sublexical route. Reduced engagement in response to sublexical route processes was also found in a left inferior parietal region, presumably engaged by attentional processes, and in a left inferior frontal region, presumably engaged by phonological processes. In contrast to this reduced engagement of the optimal left hemisphere reading network (ventral OT, inferior parietal, inferior frontal), our dyslexic readers exhibited increased engagement of visual occipital regions and of regions presumably engaged by silent articulatory processes (premotor/motor cortex and subcortical caudate and putamen).
doi:10.1016/j.cortex.2010.06.004
PMCID: PMC3073233  PMID: 20650450
6.  A Dual-Route Perspective on Brain Activation in Response to Visual Words: Evidence for a Length by Lexicality Interaction in the Visual Word Form Area (VWFA) 
NeuroImage  2009;49(3):2649-2661.
Based on our previous work, we expected the Visual Word Form Area (VWFA) in the left ventral visual pathway to be engaged by both whole-word recognition and by serial sublexical coding of letter strings. To examine this double function, a phonological lexical decision task (i.e., “Does xxx sound like an existing word?”) presented short and long letter strings of words, pseudohomophones, and pseudowords (e.g., Taxi, Taksi and Tazi). Main findings were that the length effect for words was limited to occipital regions and absent in the VWFA. In contrast, a marked length effect for pseudowords was found throughout the ventral visual pathway including the VWFA, as well as in regions presumably engaged by visual attention and silent-articulatory processes. The length by lexicality interaction on brain activation corresponds to well-established behavioral findings of a length by lexicality interaction on naming latencies and speaks for the engagement of the VWFA by both lexical and sublexical processes.
doi:10.1016/j.neuroimage.2009.10.082
PMCID: PMC2989181  PMID: 19896538
7.  On the functional neuroanatomy of visual word processing: Effects of case and letter deviance 
Journal of cognitive neuroscience  2009;21(2):222-229.
This fMRI study contrasted case-deviant and letter-deviant forms with familiar forms of the same phonological words (e.g., TaXi and Taksi vs. Taxi) and found, that both types of deviance led to increased activation in a left occipitotemporal region corresponding to the Visual Word Form Area. Case-deviant items, in addition, led to increased activation in a right occipitotemporal region and in a left occipital and a left posterior occipitotemporal region, possibly reflecting the increased demands on letter form coding. For letter-deviant items, in addition to the increased left occipitotemporal activation, a main finding was increased activation primarily in extended left frontal regions, possibly reflecting sublexically mediated access to word phonology. These findings are consistent with general features of cognitive dual-route models of visual word processing. Furthermore, they add support to the main feature of Dehaene et al.’s (2005) neural model of early stages of visual word processing . However, the increased activation found for case-deviant items in the VWFA cannot be immediately reconciled with the assumption of completely abstract case-independent orthographic word codes in the VWFA.
doi:10.1162/jocn.2009.21002
PMCID: PMC2976854  PMID: 18476755
Functional MRI; visual word recognition; occipitotemporal cortex; visual word form area; orthographic processing
8.  A dual-route perspective on poor reading in a regular orthography: An fMRI study 
This study examined functional brain abnormalities in dyslexic German readers who – due to the regularity of German in the reading direction – do not exhibit the reading accuracy problem of English dyslexic readers, but suffer primarily from a reading speed problem. The in-scanner task required phonological lexical decisions (i.e., Does xxx sound like an existing word?) and presented familiar and unfamiliar letter strings of existing phonological words (e.g., Taxi-Taksi) together with nonwords (e.g., Tazi). Dyslexic readers exhibited the same response latency pattern (words < pseudohomophones < nonwords) as nonimpaired readers, but latencies to all item types were much prolonged. The imaging results were suggestive for a different neural organization of reading processes in dyslexic readers. Specifically, dyslexic readers, in response to lexical route processes, exhibited underactivation in a left ventral occipitotemporal (OT) region which presumably is engaged by visual-orthographic whole word recognition. This region was also insensitive to the increased visual-orthographic processing demands of the sublexical route. Reduced engagement in response to sublexical route processes was also found in a left inferior parietal region, presumably engaged by attentional processes, and in a left inferior frontal region, presumably engaged by phonological processes. In contrast to this reduced engagement of the optimal left hemisphere reading network (ventral OT, inferior parietal, inferior frontal), our dyslexic readers exhibited increased engagement of visual occipital regions and of regions presumably engaged by silent articulatory processes (premotor/motor cortex and subcortical caudate and putamen).
doi:10.1016/j.cortex.2010.06.004
PMCID: PMC3073233  PMID: 20650450
Developmental dyslexia; fMRI; Reading; Phonological lexical decision; Dual-route
9.  A Common Left Occipito-Temporal Dysfunction in Developmental Dyslexia and Acquired Letter-By-Letter Reading? 
PLoS ONE  2010;5(8):e12073.
Background
We used fMRI to examine functional brain abnormalities of German-speaking dyslexics who suffer from slow effortful reading but not from a reading accuracy problem. Similar to acquired cases of letter-by-letter reading, the developmental cases exhibited an abnormal strong effect of length (i.e., number of letters) on response time for words and pseudowords.
Results
Corresponding to lesions of left occipito-temporal (OT) regions in acquired cases, we found a dysfunction of this region in our developmental cases who failed to exhibit responsiveness of left OT regions to the length of words and pseudowords. This abnormality in the left OT cortex was accompanied by absent responsiveness to increased sublexical reading demands in phonological inferior frontal gyrus (IFG) regions. Interestingly, there was no abnormality in the left superior temporal cortex which—corresponding to the onological deficit explanation—is considered to be the prime locus of the reading difficulties of developmental dyslexia cases.
Conclusions
The present functional imaging results suggest that developmental dyslexia similar to acquired letter-by-letter reading is due to a primary dysfunction of left OT regions.
doi:10.1371/journal.pone.0012073
PMCID: PMC2920311  PMID: 20711448

Results 1-9 (9)