Richard J S Wise

WWF United Kingdom, Londinium, England, United Kingdom

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Publications (111)805.63 Total impact

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    ABSTRACT: The estimated prevalence of aphasia in the UK and the USA is 250 000 and 1 000 000, respectively. The commonest aetiology is stroke. The impairment may improve with behavioural therapy, and trials using cortical stimulation or pharmacotherapy are undergoing proof-of-principle investigation, but with mixed results. Aphasia is a heterogeneous syndrome, and the simple classifications according to the Broca-Wernicke-Lichtheim model inadequately describe the diverse communication difficulties with which patients may present. Greater knowledge of how intact neural networks promote recovery after aphasic stroke, either spontaneously or in response to interventions, will result in clearer hypotheses about how to improve the treatment of aphasia. Twenty-five years ago, a pioneering study on healthy participants heralded the introduction of functional neuroimaging to the study of mechanisms of recovery from aphasia. Over the ensuing decades, such studies have been interpreted as supporting one of three hypotheses, which are not mutually exclusive. The first two predate the introduction of functional neuroimaging: that recovery is the consequence of the reconstitution of domain-specific language systems in tissue around the lesion (the 'perilesional' hypothesis), or by homotopic cortex in the contralateral hemisphere (the 'laterality-shift' hypothesis). The third is that loss of transcallosal inhibition to contralateral homotopic cortex hinders recovery (the 'disinhibition' hypothesis). These different hypotheses at times give conflicting views about rehabilitative intervention; for example, should one attempt to activate or inhibit a contralateral homotopic region with cortical stimulation techniques to promote recovery? This review proposes that although the functional imaging data are statistically valid in most cases, their interpretation has often favoured one explanation while ignoring plausible alternatives. In our view, this is particularly evident when recovery is attributed to activity in 'language networks' occupying sites not observed in healthy participants. In this review we will argue that much of the distribution of what has often been interpreted as language-specific activity, particularly in midline and contralateral cortical regions, is an upregulation of activity in intact domain-general systems for cognitive control and attention, responding in a task-dependent manner to the increased 'effort' when damaged downstream domain-specific language networks are impaired. We further propose that it is an inability fully to activate these systems that may result in sub optimal recovery in some patients. Interpretation of the data in terms of activity in domain-general networks affords insights into novel approaches to rehabilitation.
    Brain : a journal of neurology. 06/2014;
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    ABSTRACT: Spoken language production is a complex brain function that relies on large-scale networks. These include domain-specific networks that mediate language-specific processes, as well as domain-general networks mediating top-down and bottom-up attentional control. Language control is thought to involve a left-lateralized fronto-temporal-parietal (FTP) system. However, these regions do not always activate for language tasks and similar regions have been implicated in nonlinguistic cognitive processes. These inconsistent findings suggest that either the left FTP is involved in multidomain cognitive control or that there are multiple spatially overlapping FTP systems. We present evidence from an fMRI study using multivariate analysis to identify spatiotemporal networks involved in spoken language production in humans. We compared spoken language production (Speech) with multiple baselines, counting (Count), nonverbal decision (Decision), and "rest," to pull apart the multiple partially overlapping networks that are involved in speech production. A left-lateralized FTP network was activated during Speech and deactivated during Count and nonverbal Decision trials, implicating it in cognitive control specific to sentential spoken language production. A mirror right-lateralized FTP network was activated in the Count and Decision trials, but not Speech. Importantly, a second overlapping left FTP network showed relative deactivation in Speech. These three networks, with distinct time courses, overlapped in the left parietal lobe. Contrary to the standard model of the left FTP as being dominant for speech, we revealed a more complex pattern within the left FTP, including at least two left FTP networks with competing functional roles, only one of which was activated in speech production.
    The Journal of neuroscience : the official journal of the Society for Neuroscience. 06/2014; 34(26):8728-40.
  • Novraj S Dhanjal, Richard J S Wise
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    ABSTRACT: Objective: Episodic memory retrieval is reliant upon cognitive control systems, of which two have been identified with functional neuroimaging: a cingulo-opercular 'salience network' (SN), and a fronto-parietal 'executive network' (EN). In Alzheimer's disease (AD), pathology is distributed throughout higher-order cortices. The hypotheses were that this fronto-parietal pathology, would impair activity associated with verbal memory recall; and that central cholinesterase inhibition (ChI) would modulate this, improving memory recall. Methods: Functional magnetic resonance imaging was used to study normal participants and two patient groups: mild cognitive impairment (MCI) and AD. Activity within EN and SN was observed during free recall of previously heard sentences, and related to measures of recall accuracy. Results: In normals, trials with reduced recall were associated with greater activity in both SN and EN. Better recall was associated with greater activity in medial regions of the default mode network. By comparison, AD patients showed attenuated responses in both SN and EN compared with either controls or MCI patients, even after recall performance was matched between groups. Following ChI, AD patients showed no modulation of activity within SN, but increased activity within EN. There was also enhanced activity within regions associated with episodic and semantic memory, during less successful recall, requiring greater cognitive control. Interpretation: The results indicate that in AD, impaired responses of cognitive control networks during verbal memory recall are partly responsible for reduced recall performance. One action of symptom-modifying treatment is partially to reverse the abnormal function of fronto-parietal cognitive control and temporal lobe memory networks. ANN NEUROL 2014. © 2014 American Neurological Association.
    Annals of Neurology 06/2014; · 11.19 Impact Factor
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    ABSTRACT: Research on mammals predicts that the anterior striatum is a central component of human motor learning. However, as vocalizations in most mammals are innate, much of the neurobiology of human vocal learning has been inferred from studies on songbirds. Essential for song learning is a pathway, the homologue of mammalian cortical-basal ganglia 'loops', which includes the avian striatum. The present functional magnetic resonance imaging (fMRI) study investigated adult human vocal learning, a skill that persists throughout life, albeit imperfectly as late-acquired languages are spoken with an accent. Monolingual adult participants were scanned while repeating novel non-native words. After training on the pronunciation of half the words for one week, they underwent a second scan. During scanning there was no external feedback on performance. Activity declined sharply in left and right anterior striatum, both within and between scanning sessions, and this change was independent of training and performance. This indicates that adult speakers rapidly adapt to the novel articulatory movements, possibly by using motor sequences from their native speech to approximate those required for the novel speech sounds. Improved accuracy correlated only with activity in motor-sensory perisylvian cortex. We propose that future studies on vocal learning, using different behavioral and pharmacological manipulations, will provide insights into adult striatal plasticity and its potential for modification in both educational and clinical contexts.
    Journal of Neurophysiology 05/2014; · 3.30 Impact Factor
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    ABSTRACT: Understanding how dynamic changes in brain activity control behavior is a major challenge of cognitive neuroscience. Here, we consider the brain as a complex dynamic system and define two measures of brain dynamics: the synchrony of brain activity, measured by the spatial coherence of the BOLD signal across regions of the brain; and metastability, which we define as the extent to which synchrony varies over time. We investigate the relationship among brain network activity, metastability, and cognitive state in humans, testing the hypothesis that global metastability is "tuned" by network interactions. We study the following two conditions: (1) an attentionally demanding choice reaction time task (CRT); and (2) an unconstrained "rest" state. Functional MRI demonstrated increased synchrony, and decreased metastability was associated with increased activity within the frontoparietal control/dorsal attention network (FPCN/DAN) activity and decreased default mode network (DMN) activity during the CRT compared with rest. Using a computational model of neural dynamics that is constrained by white matter structure to test whether simulated changes in FPCN/DAN and DMN activity produce similar effects, we demonstate that activation of the FPCN/DAN increases global synchrony and decreases metastability. DMN activation had the opposite effects. These results suggest that the balance of activity in the FPCN/DAN and DMN might control global metastability, providing a mechanistic explanation of how attentional state is shifted between an unfocused/exploratory mode characterized by high metastability, and a focused/constrained mode characterized by low metastability.
    Journal of Neuroscience 01/2014; 34(2):451-461. · 6.91 Impact Factor
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    Richard J S Wise, Rodrigo M Braga
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    ABSTRACT: Literacy implies convergence, with heard and read narratives directed from separate sensory pathways towards common linguistic and semantic brain systems - very 'bottom-up'. But we also actively infer, relate, remember, attend and, above all, imagine - very 'top-down'. We comment on an interesting investigation of the listening and reading brain.
    Trends in Cognitive Sciences 11/2013; · 16.01 Impact Factor
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    ABSTRACT: Aphasic deficits are usually only interpreted in terms of domain-specific language processes. However, effective human communication and tests that probe this complex cognitive skill are also dependent on domain-general processes. In the clinical context, it is a pragmatic observation that impaired attention and executive functions interfere with the rehabilitation of aphasia. One system that is important in cognitive control is the salience network, which includes dorsal anterior cingulate cortex and adjacent cortex in the superior frontal gyrus (midline frontal cortex). This functional imaging study assessed domain-general activity in the midline frontal cortex, which was remote from the infarct, in relation to performance on a standard test of spoken language in 16 chronic aphasic patients both before and after a rehabilitation programme. During scanning, participants heard simple sentences, with each listening trial followed immediately by a trial in which they repeated back the previous sentence. Listening to sentences in the context of a listen-repeat task was expected to activate regions involved in both language-specific processes (speech perception and comprehension, verbal working memory and pre-articulatory rehearsal) and a number of task-specific processes (including attention to utterances and attempts to overcome pre-response conflict and decision uncertainty during impaired speech perception). To visualize the same system in healthy participants, sentences were presented to them as three-channel noise-vocoded speech, thereby impairing speech perception and assessing whether this evokes domain general cognitive systems. As expected, contrasting the more difficult task of perceiving and preparing to repeat noise-vocoded speech with the same task on clear speech demonstrated increased activity in the midline frontal cortex in the healthy participants. The same region was activated in the aphasic patients as they listened to standard (undistorted) sentences. Using a region of interest defined from the data on the healthy participants, data from the midline frontal cortex was obtained from the patients. Across the group and across different scanning sessions, activity correlated significantly with the patients' communicative abilities. This correlation was not influenced by the sizes of the lesion or the patients' chronological ages. This is the first study that has directly correlated activity in a domain general system, specifically the salience network, with residual language performance in post-stroke aphasia. It provides direct evidence in support of the clinical intuition that domain-general cognitive control is an essential factor contributing to the potential for recovery from aphasic stroke.
    Brain 10/2013; · 9.92 Impact Factor
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    ABSTRACT: Intrinsic connectivity networks (ICNs), such as the default mode, frontoparietal control, and salience networks, provide a useful large-scale description of the functional architecture of the brain. Although ICNs are functionally specialized, the information that they process needs to be integrated for coherent cognition, perception, and behavior. A region capable of performing this integration might be expected to contain traces, or "echoes," of the neural signals from multiple ICNs. Here, using fMRI in humans, we show the existence of specific "transmodal" regions containing echoes of multiple ICNs. These regions include core nodes of the default mode network, as well as multimodal association regions of the temporoparietal and temporo-occipito-parietal junction, right middle frontal gyrus, and dorsal anterior cingulate cortex. In contrast, "unimodal" regions such as the primary sensory and motor cortices show a much more singular pattern of activity, containing traces of few or even single ICNs. The presence of ICN echoes might explain how transmodal regions are involved in multiple different cognitive states. Our results suggest that these transmodal regions have a particular local spatial organization containing topographic maps that relate to multiple ICNs. This makes transmodal regions uniquely placed to be able to mediate the cross talk between the brain's functional networks through local modulation of adjacent regions that communicate with different ICNs.
    Journal of Neuroscience 08/2013; 33(35):14031-9. · 6.91 Impact Factor
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    ABSTRACT: This study tested the efficacy of audio-visual reading training in nine patients with pure alexia, an acquired reading disorder caused by damage to the left ventral occipitotemporal cortex. As well as testing the therapy's impact on reading speed, we investigated the functional reorganization underlying therapy-induced behavioural changes using magnetoencephalography. Reading ability was tested twice before training (t1 and t2) and twice after completion of the 6-week training period (t3 and t4). At t3 there was a significant improvement in word reading speed and reduction of the word length effect for trained words only. Magnetoencephalography at t3 demonstrated significant differences in reading network connectivity for trained and untrained words. The training effects were supported by increased bidirectional connectivity between the left occipital and ventral occipitotemporal perilesional cortex, and increased feedback connectivity from the left inferior frontal gyrus. Conversely, connection strengths between right hemisphere regions became weaker after training.
    Brain 08/2013; 136(Pt 8):2579-91. · 9.92 Impact Factor
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    ABSTRACT: Modern neuroimaging techniques have advanced our understanding of the distributed anatomy of speech production, beyond that inferred from clinico-pathological correlations. However, much remains unknown about functional interactions between anatomically distinct components of this speech production network. One reason for this is the need to separate spatially overlapping neural signals supporting diverse cortical functions. We took three separate human functional magnetic resonance imaging (fMRI) datasets (two speech production, one "rest"). In each we decomposed the neural activity within the left posterior perisylvian speech region into discrete components. This decomposition robustly identified two overlapping spatio-temporal components, one centered on the left posterior superior temporal gyrus (pSTG), the other on the adjacent ventral anterior parietal lobe (vAPL). The pSTG was functionally connected with bilateral superior temporal and inferior frontal regions, whereas the vAPL was connected with other parietal regions, lateral and medial. Surprisingly, the components displayed spatial anti-correlation, in which the negative functional connectivity of each component overlapped with the other component's positive functional connectivity, suggesting that these two systems operate separately and possibly in competition. The speech tasks reliably modulated activity in both pSTG and vAPL suggesting they are involved in speech production, but their activity patterns dissociate in response to different speech demands. These components were also identified in subjects at "rest" and not engaged in overt speech production. These findings indicate that the neural architecture underlying speech production involves parallel distinct components that converge within posterior peri-sylvian cortex, explaining, in part, why this region is so important for speech production. Hum Brain Mapp, 2013. © 2013 Wiley Periodicals, Inc.
    Human Brain Mapping 05/2013; · 6.88 Impact Factor
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    ABSTRACT: An anterior pathway, concerned with extracting meaning from sound, has been identified in nonhuman primates. An analogous pathway has been suggested in humans, but controversy exists concerning the degree of lateralization and the precise location where responses to intelligible speech emerge. We have demonstrated that the left anterior superior temporal sulcus (STS) responds preferentially to intelligible speech (Scott SK, Blank CC, Rosen S, Wise RJS. 2000. Identification of a pathway for intelligible speech in the left temporal lobe. Brain. 123:2400-2406.). A functional magnetic resonance imaging study in Cerebral Cortex used equivalent stimuli and univariate and multivariate analyses to argue for the greater importance of bilateral posterior when compared with the left anterior STS in responding to intelligible speech (Okada K, Rong F, Venezia J, Matchin W, Hsieh IH, Saberi K, Serences JT,Hickok G. 2010. Hierarchical organization of human auditory cortex: evidence from acoustic invariance in the response to intelligible speech. 20: 2486-2495.). Here, we also replicate our original study, demonstrating that the left anterior STS exhibits the strongest univariate response and, in decoding using the bilateral temporal cortex, contains the most informative voxels showing an increased response to intelligible speech. In contrast, in classifications using local "searchlights" and a whole brain analysis, we find greater classification accuracy in posterior rather than anterior temporal regions. Thus, we show that the precise nature of the multivariate analysis used will emphasize different response profiles associated with complex sound to speech processing.
    Cerebral Cortex 04/2013; · 8.31 Impact Factor
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    ABSTRACT: A central question for cognitive neuroscience is whether there is a single neural system controlling the allocation of attention. A dorsal frontoparietal network of brain regions is often proposed as a mediator of top-down attention to all sensory inputs. We used functional magnetic resonance imaging in humans to show that the cortical networks supporting top-down attention are in fact modality-specific, with distinct superior fronto-parietal and fronto-temporal networks for visuospatial and non-spatial auditory attention respectively. In contrast, parts of the right middle and inferior frontal gyri showed a common response to attentional control regardless of modality, providing evidence that the amodal component of attention is restricted to the anterior cortex.
    NeuroImage 02/2013; · 6.25 Impact Factor
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    ABSTRACT: OBJECTIVE: Episodic memory encoding of a verbal message depends upon initial registration, which requires sustained auditory attention followed by deep semantic processing of the message. Motivated by previous data demonstrating modulation of auditory cortical activity during sustained attention to auditory stimuli, we investigated the response of the human auditory cortex during encoding of sentences to episodic memory. Subsequently, we investigated this response in patients with mild cognitive impairment (MCI) and probable Alzheimer's disease (pAD). METHODS: Using functional magnetic resonance imaging, 31 healthy participants were studied. The response in 18 MCI and 18 pAD patients was then determined, and compared to 18 matched healthy controls. Subjects heard factual sentences, and subsequent retrieval performance indicated successful registration and episodic encoding. RESULTS: The healthy subjects demonstrated that suppression of auditory cortical responses was related to greater success in encoding heard sentences; and that this was also associated with greater activity in the semantic system. In contrast, there was reduced auditory cortical suppression in patients with MCI, and absence of suppression in pAD. Administration of a central cholinesterase inhibitor (ChI) partially restored the suppression in patients with pAD, and this was associated with an improvement in verbal memory. INTERPRETATION: Verbal episodic memory impairment in AD is associated with altered auditory cortical function, reversible with a ChI. Although these results may indicate the direct influence of pathology in auditory cortex, they are also likely to indicate a partially reversible impairment of feedback from neocortical systems responsible for sustained attention and semantic processing. ANN NEUROL 2012.
    Annals of Neurology 10/2012; · 11.19 Impact Factor
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    ABSTRACT: This functional MRI study investigated the involvement of the left inferior parietal cortex (IPC) in spoken language production (Speech). Its role has been apparent in some studies but not others, and is not convincingly supported by clinical studies as they rarely include cases with lesions confined to the parietal lobe. We compared Speech with non-communicative repetitive tongue movements (Tongue). The data were analyzed with both univariate contrasts between conditions and probabilistic independent component analysis (ICA). The former indicated decreased activity of left IPC during Speech relative to Tongue. However, the ICA revealed a Speech component in which there was correlated activity between left IPC, frontal and temporal cortices known to be involved in language. Therefore, although net synaptic activity throughout the left IPC may not increase above baseline conditions during Speech, one or more local systems within this region are involved, evidenced by the correlated activity with other language regions.
    Brain and Language 02/2012; 121(1):47-57. · 3.39 Impact Factor
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    Zarinah K Agnew, Richard J S Wise, Robert Leech
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    ABSTRACT: Mirror neurons are single cells found in macaque premotor and parietal cortices that are active during action execution and observation. In non-human primates, mirror neurons have only been found in relation to object-directed movements or communicative gestures, as non-object directed actions of the upper limb are not well characterized in non-human primates. Mirror neurons provide important evidence for motor simulation theories of cognition, sometimes referred to as the direct matching hypothesis, which propose that observed actions are mapped onto associated motor schemata in a direct and automatic manner. This study, for the first time, directly compares mirror responses, defined as the overlap between action execution and observation, during object directed and meaningless non-object directed actions. We present functional MRI data that demonstrate a clear dissociation between object directed and non-object directed actions within the human mirror system. A premotor and parietal network was preferentially active during object directed actions, whether observed or executed. Moreover, we report spatially correlated activity across multiple voxels for observation and execution of an object directed action. In contrast to predictions made by motor simulation theory, no similar activity was observed for non-object directed actions. These data demonstrate that object directed and meaningless non-object directed actions are subserved by different neuronal networks and that the human mirror response is significantly greater for object directed actions. These data have important implications for understanding the human mirror system and for simulation theories of motor cognition. Subsequent theories of motor simulation must account for these differences, possibly by acknowledging the role of experience in modulating the mirror response.
    PLoS ONE 01/2012; 7(4):e32517. · 3.53 Impact Factor
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    ABSTRACT: One functional anatomical model of reading, drawing on human neuropsychological and neuroimaging data, proposes that a region in left ventral occipitotemporal cortex (vOT) becomes, through experience, specialized for written word perception. We tested this hypothesis by presenting numbers in orthographical and digital form with two task demands, phonological and numerical. We observed a main effect of task on left vOT activity but not stimulus type, with increased activity during the phonological task that was also associated with increased activity in the left inferior frontal gyrus, a region implicated in speech production. Region-of-interest analysis confirmed that there was equal activity for orthographical and digital written forms in the left vOT during the phonological task, despite greater visual complexity of the orthographical forms. This evidence is incompatible with a predominantly feedforward model of written word recognition that proposes that the left vOT is a specialized cortical module for word recognition in literate subjects. Rather, the physiological data presented here fits better with interactive computational models of reading that propose that written word recognition emerges from bidirectional interactions between three processes: visual, phonological, and semantic. Further, the present study is in accord with others that indicate that the left vOT is a route through which nonlinguistic stimuli, perhaps high contrast two-dimensional objects in particular, gain access to a predominantly left-lateralized language and semantic system.
    Journal of Neuroscience 10/2011; 31(41):14631-8. · 6.91 Impact Factor
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    ABSTRACT: Localising activity in the human midbrain with conventional functional MRI (fMRI) is challenging because the midbrain nuclei are small and located in an area that is prone to physiological artefacts. Here we present a replicable and automated method to improve the detection and localisation of midbrain fMRI signals. We designed a visual fMRI task that was predicted would activate the superior colliculi (SC) bilaterally. A limited number of coronal slices were scanned, orientated along the long axis of the brainstem, whilst simultaneously recording cardiac and respiratory traces. A novel anatomical registration pathway was used to optimise the localisation of the small midbrain nuclei in stereotactic space. Two additional structural scans were used to improve registration between functional and structural T1-weighted images: an echo-planar image (EPI) that matched the functional data but had whole-brain coverage, and a whole-brain T2-weighted image. This pathway was compared to conventional registration pathways, and was shown to significantly improve midbrain registration. To reduce the physiological artefacts in the functional data, we estimated and removed structured noise using a modified version of a previously described physiological noise model (PNM). Whereas a conventional analysis revealed only unilateral SC activity, the PNM analysis revealed the predicted bilateral activity. We demonstrate that these methods improve the measurement of a biologically plausible fMRI signal. Moreover they could be used to investigate the function of other midbrain nuclei.
    NeuroImage 08/2011; 59(2):1230-8. · 6.25 Impact Factor
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    ABSTRACT: A foreign language (L2) learned after childhood results in an accent. This functional neuroimaging study investigated speech in L2 as a sensory-motor skill. The hypothesis was that there would be an altered response in auditory and somatosensory association cortex, specifically the planum temporale and parietal operculum, respectively, when speaking in L2 relative to L1, independent of rate of speaking. These regions were selected for three reasons. First, an influential computational model proposes that these cortices integrate predictive feedforward and postarticulatory sensory feedback signals during articulation. Second, these adjacent regions (known as Spt) have been identified as a "sensory-motor interface" for speech production. Third, probabilistic anatomical atlases exist for these regions, to ensure the analyses are confined to sensory-motor differences between L2 and L1. The study used functional magnetic resonance imaging (fMRI), and participants produced connected overt speech. The first hypothesis was that there would be greater activity in the planum temporale and the parietal operculum when subjects spoke in L2 compared with L1, one interpretation being that there is less efficient postarticulatory sensory monitoring when speaking in the less familiar L2. The second hypothesis was that this effect would be observed in both cerebral hemispheres. Although Spt is considered to be left-lateralized, this is based on studies of covert speech, whereas overt speech is accompanied by sensory feedback to bilateral auditory and somatosensory cortices. Both hypotheses were confirmed by the results. These findings provide the basis for future investigations of sensory-motor aspects of language learning using serial fMRI studies.
    Journal of Neurophysiology 05/2011; 106(1):470-8. · 3.30 Impact Factor
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    Zoe Victoria Joan Woodhead, Richard James Surtees Wise, Marty Sereno, Robert Leech
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    ABSTRACT: Different cortical regions within the ventral occipitotemporal junction have been reported to show preferential responses to particular objects. Thus, it is argued that there is evidence for a left-lateralized visual word form area and a right-lateralized fusiform face area, but the unique specialization of these areas remains controversial. Words are characterized by greater power in the high spatial frequency (SF) range, whereas faces comprise a broader range of high and low frequencies. We investigated how these high-order visual association areas respond to simple sine-wave gratings that varied in SF. Using functional magnetic resonance imaging, we demonstrated lateralization of activity that was concordant with the low-level visual property of words and faces; left occipitotemporal cortex is more strongly activated by high than by low SF gratings, whereas the right occipitotemporal cortex responded more to low than high spatial frequencies. Therefore, the SF of a visual stimulus may bias the lateralization of processing irrespective of its higher order properties.
    Cerebral Cortex 03/2011; 21(10):2307-12. · 8.31 Impact Factor
  • Brain and Language 03/2011; 116(3):103-4. · 3.39 Impact Factor

Publication Stats

7k Citations
805.63 Total Impact Points

Institutions

  • 2014
    • WWF United Kingdom
      Londinium, England, United Kingdom
  • 2003–2014
    • Imperial College London
      • • Division of Experimental Medicine
      • • Faculty of Medicine
      Londinium, England, United Kingdom
  • 2003–2012
    • University College London
      • • Institute of Cognitive Neuroscience
      • • Division of Psychology and Language Sciences
      London, ENG, United Kingdom
  • 2001–2011
    • MRC Clinical Sciences Centre
      London Borough of Harrow, England, United Kingdom
  • 2006
    • Oxford University Hospitals NHS Trust
      • Department of Clinical Neurology
      Oxford, England, United Kingdom
  • 2001–2002
    • University of Oxford
      • Nuffield Department of Clinical Neurosciences
      Oxford, ENG, United Kingdom