Cortical localisation of the visual and auditory word form areas: a reconsideration of the evidence

In this paper we examine the evidence for human brain areas dedicated to visual or auditory word form processing by comparing cortical activation for auditory word repetition, reading, picture naming, and environmental sound naming. Both reading and auditory word repetition activated left lateralised regions in the frontal operculum (Broca's area), posterior superior temporal gyrus (Wernicke's area), posterior inferior temporal cortex, and a region in the mid superior temporal sulcus relative to baseline conditions that controlled for sensory input and motor output processing. In addition, auditory word repetition increased activation in a lateral region of the left mid superior temporal gyrus but critically, this area is not specific to auditory word processing, it is also activated in response to environmental sounds. There were no reading specific activations, even in the areas previously claimed as visual word form areas: activations were either common to reading and auditory word repetition or common to reading and picture naming. We conclude that there is no current evidence for cortical sites dedicated to visual or auditory word form processing.     

Lesion analysis of the brain areas involved in language comprehension

The cortical regions of the brain traditionally associated with the comprehension of language are Wernicke's area and Broca's area. However, recent evidence suggests that other brain regions might also be involved in this complex process. This paper describes the opportunity to evaluate a large number of brain-injured patients to determine which lesioned brain areas might affect language comprehension. Sixty-four chronic left hemisphere stroke patients were evaluated on 11 subtests of the Curtiss-Yamada Comprehensive Language Evaluation - Receptive (CYCLE-R; Curtiss, S., & Yamada, J. (1988). Curtiss-Yamada Comprehensive Language Evaluation. Unpublished test, UCLA). Eight right hemisphere stroke patients and 15 neurologically normal older controls also participated. Patients were required to select a single line drawing from an array of three or four choices that best depicted the content of an auditorily-presented sentence. Patients' lesions obtained from structural neuroimaging were reconstructed onto templates and entered into a voxel-based lesion-symptom mapping (VLSM; Bates, E., Wilson, S., Saygin, A. P., Dick, F., Sereno, M., Knight, R. T., & Dronkers, N. F. (2003). Voxel-based lesion-symptom mapping. Nature Neuroscience, 6(5), 448-450.) analysis along with the behavioral data. VLSM is a brain-behavior mapping technique that evaluates the relationships between areas of injury and behavioral performance in all patients on a voxel-by-voxel basis, similar to the analysis of functional neuroimaging data. Results indicated that lesions to five left hemisphere brain regions affected performance on the CYCLE-R, including the posterior middle temporal gyrus and underlying white matter, the anterior superior temporal gyrus, the superior temporal sulcus and angular gyrus, mid-frontal cortex in Brodmann's area 46, and Brodmann's area 47 of the inferior frontal gyrus. Lesions to Broca's and Wernicke's areas were not found to significantly alter language comprehension on this particular measure. Further analysis suggested that the middle temporal gyrus may be more important for comprehension at the word level, while the other regions may play a greater role at the level of the sentence. These results are consistent with those seen in recent functional neuroimaging studies and offer complementary data in the effort to understand the brain areas underlying language comprehension.     

Evidence for right hemisphere phonology in a backward masking task

The extent to which orthographic and phonological processes are available during the initial moments of word recognition within each hemisphere is under specified, particularly for the right hemisphere. Few studies have investigated whether each hemisphere uses orthography and phonology under constraints that restrict the viewing time of words and reduce overt phonological demands. The current study used backward masking in the divided visual field paradigm to explore hemisphere differences in the availability of orthographic and phonological word recognition processes. A 20 ms and 60 ms SOA were used to track the time course of how these processes develop during pre-lexical moments of word recognition. Nonword masks varied in similarity to the target words such that there were four types: orthographically and phonologically similar, orthographically but not phonologically similar, phonologically but not orthographically similar and unrelated. The results showed the left hemisphere has access to both orthography and phonology early in the word recognition process. With more time to process the stimulus, the left hemisphere is able to use phonology which benefits word recognition to a larger extent than orthography. The right hemisphere also demonstrates access to both orthography and phonology in the initial moments of word recognition, however, orthographic similarity improves word recognition to a greater extent than phonological similarity.     

Vowel perception: A neuroradiological localization of the perception of vowels in the human cortex

Whether the perception of vowels takes place in the right or left cerebral hemisphere, or is dependent on bilateral cortical processes, is of importance in our treatment of patients with language deficiencies. To investigate this problem a phonetic test with a series of vowellike stimuli was administered to 68 right-handed adult patients with cerebral lesions, mainly of vascular origin. The results were compared to the results in a control group of 19 speech therapists. Second, a neuroradiologic method was developed in order to visualize the anatomic site of Wernicke's area in the left hemisphere on CT scans. This method formed the basis for an evaluation of the extent and localization of the patients' lesions in Wernicke's area. Of 46 patients with lesions in the left hemisphere, 19 had no perceptual disturbances and 27 had severe perceptual disturbances with lesions predominantly located in Wernicke's area. Twenty-two patients with lesions in the corresponding area in the right hemisphere showed no perceptual disturbances. The results of this investigation appear to indicate that the perception of vowels in right-handed persons is unilaterally located in Wernicke's area in the left hemisphere.     

Neuromagnetic correlates of intra- and extra-dimensional set-shifting

Set-shifting is essential to cognitive flexibility and relies on frontal lobe function. Previous studies have mostly focused on feedback processes following shifting rather than set-shifting itself. We designed an MEG paradigm without feedback to directly investigate the neural correlates of set-shifting. Adults (n = 16) matched one of two coloured images with a third stimulus, the target, by either the colour or shape dimension of the target. Half of the shift trials involved colour-to-colour or shape-to-shape (intra-dimensional: ID) shifting and the other half involved colour-to-shape or shape-to-colour (extra-dimensional: ED) shifting. MEG was continuously recorded on a 151 channel CTF system. We used beamforming to analyze responses to the first (shift) and the third (repeat) trials in each set. These trials were contrasted separately for ID and ED sets. Shift versus repeat trials showed larger MEG activations for intra-dimensional shifting in the right inferior frontal gyrus (BA 47), left medial frontal gyrus (BA 10) and right superior frontal gyrus (BA 9) as early as 100 ms, and in left middle frontal gyrus (BA 11) between 250–500 ms. Activations related to extra-dimensional shifting were detected in left inferior frontal gyrus (BA 44), left middle frontal gyrus (BA 11), and right middle frontal gyrus (BA 46) between 100 ms and 350 ms, followed by superior frontal gyrus (BA 8/BA 10) between 250–500 ms. Intra-dimensional and extra-dimensional shifting also activated bilateral and right parietal areas, respectively. This study establishes the location and timing of frontal and parietal activations during an intra-dimensional versus extra-dimensional shifting task.     

Dissociation and association of the embodied representation of tool-use verbs and hand verbs: An fMRI study

Embodied semantic theories suppose that representation of word meaning and actual sensory-motor processing are implemented in overlapping systems. According to this view, association and dissociation of different word meaning should correspond to dissociation and association of the described sensory-motor processing. Previous studies demonstrate that although tool-use actions and hand actions have overlapping neural substrates, tool-use actions show greater activations in frontal–parietal–temporal regions that are responsible for motor control and tool knowledge processing. In the present study, we examined the association and the dissociation of the semantic representation of tool-use verbs and hand action verbs. Chinese verbs describing tool-use or hand actions without tools were included, and a passive reading task was employed. All verb conditions showed common activations in areas of left middle frontal gyrus, left inferior frontal gyrus (BA 44/45) and left inferior parietal lobule relative to rest, and all conditions showed significant effects in premotor areas within the mask of hand motion effects. Contrasts between tool-use verbs and hand verbs demonstrated that tool verbs elicited stronger activity in left superior parietal lobule, left middle frontal gyrus and left posterior middle temporal gyrus. Additionally, psychophysiological interaction analyses demonstrated that tool verbs indicated greater connectivity among these regions. These results suggest that the brain regions involved in tool-use action processing also play more important roles in tool-use verb processing and that similar systems may be responsible for word meaning representation and actual sensory-motor processing.     

Bilateral capacity for speech sound processing in auditory comprehension: evidence from Wada procedures

Data from lesion studies suggest that the ability to perceive speech sounds, as measured by auditory comprehension tasks, is supported by temporal lobe systems in both the left and right hemisphere. For example, patients with left temporal lobe damage and auditory comprehension deficits (i.e., Wernicke's aphasics), nonetheless comprehend isolated words better than one would expect if their speech perception system had been largely destroyed (70-80% accuracy). Further, when comprehension fails in such patients their errors are more often semantically-based, than-phonemically based. The question addressed by the present study is whether this ability of the right hemisphere to process speech sounds is a result of plastic reorganization following chronic left hemisphere damage, or whether the ability exists in undamaged language systems. We sought to test these possibilities by studying auditory comprehension in acute left versus right hemisphere deactivation during Wada procedures. A series of 20 patients undergoing clinically indicated Wada procedures were asked to listen to an auditorily presented stimulus word, and then point to its matching picture on a card that contained the target picture, a semantic foil, a phonemic foil, and an unrelated foil. This task was performed under three conditions, baseline, during left carotid injection of sodium amytal, and during right carotid injection of sodium amytal. Overall, left hemisphere injection led to a significantly higher error rate than right hemisphere injection. However, consistent with lesion work, the majority (75%) of these errors were semantic in nature. These findings suggest that auditory comprehension deficits are predominantly semantic in nature, even following acute left hemisphere disruption. This, in turn, supports the hypothesis that the right hemisphere is capable of speech sound processing in the intact brain.     

Structural influences on the resolution of lexical ambiguity: an analysis of hemispheric asymmetries

Structural influences on lexical ambiguity resolution in the two cerebral hemispheres was investigated using a divided visual field procedure. Participants were presented with auditory Wh- sentences containing an ambiguous word, where the grammatical role of the word was apparent only at a sentence-final verb (e.g., "Which BANK did the woman see?"). Following a sentence, either immediately or after 600 ms, a target word was presented in either the right or left visual field. Targets were related to the ambiguous word's dominant meaning (MONEY), the subordinate meaning (RIVER), or were unrelated. With left visual field presentation, priming occurred for both dominant- and subordinate-related targets at a 0 ms delay, but only for dominant-related targets at 600 ms. With left visual field presentation, priming occurred for subordinate-related targets only at both delays. The results suggest that grammatical assignment triggers the selection of meaning in the left hemisphere, whereas processing in the right hemisphere operates independently of structural analyses.     

默读汉字词的脑功能偏侧化成像研究

采用读词名的方法 ,对 12名正常大学生 ,运用功能性磁共振成像技术 (fMRI) ,研究了汉字词及假词视觉识别时的脑功能偏侧化现象。实验结果表明 ,汉字词激活左下额区 (BA4 5 )以及右颞叶 (BA2 1、2 2 )、右枕叶 (BA18) ,而假词除额区外表现出广泛的激活。此结果显示除左脑半球与汉字词加工密切相关外 ,右脑半球在汉字词加工中有一定参与但其确切作用有待进一步研究     

Inter-hemispheric remapping between arm proprioception and vision of the hand is disrupted by single pulse TMS on the left parietal cortex

Parietal cortical areas are involved in sensori-motor transformations for their respective contralateral hemifield/body. When arms of the subjects are crossed while their gaze is fixed straight ahead, vision of the hand is processed by the hemisphere ipsilateral to the arm position and proprioception of the arm by the contralateral hemisphere. It induces interhemispheric transfer and remapping. Our objective was to investigate whether a single pulse TMS applied to the left parietal cortical area would disturb interhemispheric remapping in a similar case, and would increase a simple reaction time (RT) with respect to a control single pulse TMS applied to the frontal cortical area. Two LED were superimposed and located in front of the subjects on the saggital axis. Subjects were asked to carefully fixate on these LED during each trial. The lighting of the red LED was used as a warning signal. Following the green one was illuminated after a variable delay and served as a go-signal. The hand for the response was determined before the start of each trial. TMS was applied to the left parietal, the left frontal cortical areas, or not applied to the subject. Results revealed that: (1) Irrespective of its location, single pulse TMS induced a non-specific effect similar to a startle reflex and reduced RT substantially (15 ms on average) with respect to a control condition without TMS (mean value = 153 ms). (2) Irrespective of TMS, RT were shorter when the right or the left hand was positioned in the right visual hemi-field (i.e. normal and crossed positions respectively). (3) Finally, RT increased when single pulse TMS was applied to the left parietal area and when hands were crossed irrespective of which hand was used. We concluded that interhemispheric sensori-motor remapping was disrupted by a single pulse TMS that was applied to the left parietal cortex. This effect was also combined with some visual attention directed towards the hand located on the right visual hemi-field.     

幽默加工中不一致探测与解决的脑机制分离

本fMRI研究使用“文字背景—漫画图片”实验范式,设计了三种实验条件（可笑、不可笑和无关）,试图对幽默加工中不一致探测与解决的脑机制进行分离。可笑与不可笑条件对比的结果与前人研究一致,证明了新实验范式的有效性。其次,无关与不可笑条件比较,右侧脑岛激活,说明脑岛可能与不一致探测有关。综合分析发现,脑岛与杏仁核可能共同参与了不一致探测加工。此外,结果还第一次区分了幽默加工中不一致解决与一般问题解决的脑区。     

Using fMRI to study recovery from acquired dysphasia

We have used functional magnetic resonance imaging (fMRI) to characterize brain activations associated with two distinct language tasks performed by a 28-year-old woman after partial recovery from dysphasia due to a left frontal hemispheric ischemic stroke. MRI showed that her ischemic lesion extended posteriorly from the left inferior frontal to the perisylvian cortex. fMRI scans of both language tasks revealed substantial differences in activation pattern relative to controls. The nature of this difference was task-specific. During performance of a verbal semantic decision task, the patient, in contrast to controls, activated a network of brain areas that excluded the inferior frontal gyrus (in either hemisphere). A second task involving rhyme judgment was designed to place a heavier cognitive load on language production processes and activated the left inferior frontal gyrus (Broca's area) strongly in normal controls. During this task, the most prominent frontal activation in the patient occurred in the right homologue of Broca's area. Subsequent analysis of this data by methods able to deal with responses of changing amplitude revealed additional, less sustained recruitment by the patient of cortex adjacent to the infarct in the region inferior to Broca's area during rhyming. These results suggest that in addition to changes in cognitive strategy, recovery from dysphasia could be mediated by both the preservation of neuronal networks in and around the infarct and the use of homologous regions in the contralateral hemisphere.     

Brain activation during semantic processing in autism spectrum disorders via functional magnetic resonance imaging

Language and communication deficits are core features of autism spectrum disorders (ASD), even in high-functioning adults with ASD. This study investigated brain activation patterns using functional magnetic resonance imaging in right-handed adult males with ASD and a control group, matched on age, handedness, and verbal IQ. Semantic processing in the controls produced robust activation in Broca's area (left inferior frontal gyrus) and in superior medial frontal gyrus and right cerebellum. The ASD group had substantially reduced Broca's activation, but increased left temporal (Wernicke's) activation. Furthermore, the ASD group showed diminished activation differences between concrete and abstract words, consistent with behavioral studies. The current study suggests Broca's area is a region of abnormal neurodevelopment in ASD, which may be linked with semantic and related language deficits frequently observed in ASD.     

Contextual modulation of N400 amplitude to lexically ambiguous words

Though much is known about the N400 component, an event-related EEG potential that is sensitive to semantic manipulations, it is unclear whether modulations of N400 amplitude reflect automatic processing, controlled processing, or both. We examined this issue using a semantic judgment task that manipulated local and global contextual cues. Word triplets (prime-noun-target, e.g., finance-bank-money) were sequentially presented on a computer screen (500 ms duration, 1000 ms stimulus onset asynchrony), in which the second word was a homograph. The first word (prime) created a neutral-, dominant-meaning- or subordinate-meaning-biased "global context," and the third word (target) created a dominant- or subordinate-biased "local context" that was either congruent or incongruent with the "global context" established by the first prime word. Participants were instructed to read all three words but to decide only whether the second and third words were semantically related. Event-related potentials (ERPs), specifically the N400, were recorded to the third terminal word. N400 amplitudes evoked by dominant meaning-related third words incongruent with the globally biased subordinate context (e.g., river-bridge-money) were significantly more negative than dominant endings in neutral contexts (e.g., taxi-bank-money), but not different from unrelated filler triplets. In addition, there was some indication that left hemisphere, temporal-parietal electrode sites were associated with greater N400 negativity for dominant targets in conflicting subordinate global contexts than homologous right hemisphere electrode sites, the latter of which showed significant activation to subordinate meanings in cooperating contexts. Thus, N400 amplitude was more affected by global than local context suggesting that controlled processes may take priority over automatic processes in modulating N400 amplitude, especially for left hemisphere electrode sites.     

Crossed Wernicke's aphasia: a case report

Crossed aphasia is a phenomenon in which an individual sustains a lesion in the right hemisphere (typically non-language dominant), but who exhibits an aphasic syndrome. The authors present a case study of an individual with crossed aphasia (CA) in an attempt to provide anecdotal information for four questions posed by : (a). Is CA a reversal of the normal cerebral hemisphere pattern of language function? (b). Does the presence of aphasia following a right cerebral hemisphere lesion indicate that typical right hemisphere functions (e.g., visual perception) are intact? (c). How may the aphasia's presentation differ from typical left hemisphere aphasias? And (d). is the pattern of improvement following CA similar to that of typical left hemisphere aphasias? We longitudinally examined the communicative-cognitive performance of an adult man with crossed aphasia of the Wernicke's type following a cerebrovascular accident. A 21-week follow-up evaluation indicated improvements in his language functioning from our initial evaluation, but he continued to exhibit a classic, moderately severe Wernicke's aphasia.     

Parallel but temporally displaced visual half-field metacontrast functions

Using a classic letter-ring metacontrast paradigm, left and right visual field meta-contrast functions were separately determined. The parallel U-shaped recognition functions for both half-fields were found to interact differentially with stimulus onset asynchrony, the left visual field function being displaced by 13 ms toward longer test stimulus-masking stimulus separations. This result was consistent with the hypothesis of longer processing time requirements for verbal stimuli delivered to the right than to the left hemisphere. This indicates that the neural locus (loci) responsible for left visual field verbal processing delay is (are) capable of mediating metacontrast phenomena. It was tentatively concluded that a relative processing delay within the right hemisphere underlies the differing visual half-field metacontrast interaction with stimulus onset asynchrony.     

In Search of the Self: A Positron Emission Tomography Study

Abstract—Previous work using positron emission tomography (PET) has shown that memory encoding processes are associated with preferential activation of left frontal regions of the brain, whereas retrieval processes are associated predominantly with right frontal activations. One possible reason for the asymmetry is that episodic retrieval necessarily involves reference to the self, and the self-concept may be represented (at least partially) in right frontal regions. Accordingly, the present study investigated the possibility that encoding of self-related material might also activate right frontal areas. Eight right-handed volunteers judged trait adjectives under four separate PET scan conditions: (a) relevance to self, (b) relevance to a well-known public figure, (c) social desirability, and (d) number of syllables. The results showed that self-related encoding yielded left frontal activations similar to those associated with other types of semantic encoding, but also specific activations in the right frontal lobe. It is concluded that the concept of self involves both general schematic structures and further specific components involved in episodic memory retrieval.     

Interhemispheric communication via direct connections for alternative meanings of ambiguous words

A priming experiment was used to investigate Burgess and Simpson's (1988) claim that interhemispheric cooperation plays an essential role in the interpretation of ambiguous text. In doing so, the merits of two models of interhemispheric cooperation, the homotopic inhibition theory (Cook, 1986) and the direct connections model (Collins & Coney, 1998), were examined. Priming of alternative meanings of ambiguous words was measured using homographs and their dominant (e.g., BARK-DOG) and subordinate meanings (e.g., BARK-TREE) as related pairs in a lexical decision task, with normal university students as subjects. Stimulus pairs were temporally separated by stimulus onset asynchronies (SOAs) of 180 and 350 ms and were independently projected to the left or right visual fields (LVF or RVF). At the shorter SOA, priming was restricted to LVF-RVF presentations, with homograph primes directed to the LVF equally facilitating responses to RVF targets which were associated with their dominant and subordinate meanings. This suggests that within 180 ms, a homograph projected to the right hemisphere activates a range of alternative meanings in the left hemisphere. At an SOA of 350 ms, LVF-RVF priming was obtained along with RVF-LVF and RVF-RVF priming. Evidently at this stage of processing, an ambiguous word directed to either hemisphere activates a range of alternative meanings in the contralateral hemisphere, while RVF primes also activate subordinate, but not dominant meanings in the left hemisphere. A homograph directed to the LVF did not activate dominant or subordinate meanings within the right hemisphere at either SOA. Generally, ambiguous words directed to either hemisphere activated a more extensive array of meanings in the contralateral hemisphere than in the hemisphere to which the prime was directed. This confirms the importance of interhemispheric cooperation in generating alternate meanings of ambiguous words. Strong support was found for the direct connections model (Collins & Coney, 1998), but no support for the homotopic inhibition theory (Cook, 1986).     

Recovery of single word comprehension: CT-scan correlates

9tory comprehension for single words, as measured by the BDAE Word Discrimination subtest, was studied longitudinally in a group of 37 aphasic patients. The majority of the patients had impaired performance at 1 month postonset, but recovery by Month 6 postonset was quite striking. Damage to Wernicke's area did not preclude recovery of single word comprehension to within the normal range. Only patients with very extensive left hemisphere lesions had incomplete recovery of single word comprehension.     

Hemisphere differences in conscious and unconscious word reading

Hemisphere differences in word reading were examined using explicit and implicit processing measures. In an inclusion task, which indexes both conscious (explicit) and unconscious (implicit) word reading processes, participants were briefly presented with a word in either the right or the left visual field and were asked to use this word to complete a three-letter word stem. In an exclusion task, which estimates unconscious word reading, participants completed the word stem with any word other than the prime word. Experiment 1 showed that words presented to either visual field were processed in very similar ways in both tasks, with the exception that words in the right visual field (left hemisphere) were more readily accessible for conscious report. Experiment 2 indicated that unconsciously processed words are shared between the hemispheres, as similar results were obtained when either the same or the opposite visual field received the word stem. Experiment 3 demonstrated that this sharing between hemispheres is cortically mediated by testing a split-brain patient. These results suggest that the left hemisphere advantage for word reading holds only for explicit measures; unconscious word reading is much more balanced between the hemispheres.