The lateral‐occipital and the inferior‐frontal cortex play different roles during the naming of visually presented objects

We reasoned that if an area is devoted to processing only the visual features of objects, then transcranial magnetic stimulation (TMS) applied to this area in either hemisphere would affect the naming of objects presented in contralateral but not ipsilateral space. In contrast, if an area is involved in language, then one might expect to see effects of TMS when applied over the left but not the right hemisphere, regardless whether objects are in contralateral or ipsilateral space. Our experiments reveal two important findings. First, TMS delivered to the lateral‐occipital complex (LOC), a visual‐form area, affected the naming of objects presented in contralateral but not ipsilateral space, independent of which hemisphere was stimulated. In two additional experiments, when participants named the color of objects or made judgments about the size of stimuli as shown physically on a computer screen, TMS over the contralateral LOC did not affect color naming but did affect the participants' ability to make size judgments. Second, TMS delivered to the left but not the right posterior inferior‐frontal gyrus (pIFG) affected the naming of objects irrespective of whether objects were presented in contralateral or ipsilateral space. In a separate experiment, when participants were asked to either read or categorize words, TMS over the left but not the right pIFG affected word categorization but not word reading. On the basis of these findings, we propose that when people name visually‐presented objects, LOC processes the visual form of objects while the left pIFG processes the semantics of objects. Hum Brain Mapp, 2009. © 2009 Wiley‐Liss, Inc.     

Right‐hemispheric processing of non‐linguistic word features: Implications for mapping language recovery after stroke

Verbal stimuli often induce right‐hemispheric activation in patients with aphasia after left‐hemispheric stroke. This right‐hemispheric activation is commonly attributed to functional reorganization within the language system. Yet previous evidence suggests that functional activation in right‐hemispheric homologues of classic left‐hemispheric language areas may partly be due to processing nonlinguistic perceptual features of verbal stimuli. We used functional MRI (fMRI) to clarify the role of the right hemisphere in the perception of nonlinguistic word features in healthy individuals. Participants made perceptual, semantic, or phonological decisions on the same set of auditorily and visually presented word stimuli. Perceptual decisions required judgements about stimulus‐inherent changes in font size (visual modality) or fundamental frequency contour (auditory modality). The semantic judgement required subjects to decide whether a stimulus is natural or man‐made; the phonologic decision required a decision on whether a stimulus contains two or three syllables. Compared to phonologic or semantic decision, nonlinguistic perceptual decisions resulted in a stronger right‐hemispheric activation. Specifically, the right inferior frontal gyrus (IFG), an area previously suggested to support language recovery after left‐hemispheric stroke, displayed modality‐independent activation during perceptual processing of word stimuli. Our findings indicate that activation of the right hemisphere during language tasks may, in some instances, be driven by a “nonlinguistic perceptual processing” mode that focuses on nonlinguistic word features. This raises the possibility that stronger activation of right inferior frontal areas during language tasks in aphasic patients with left‐hemispheric stroke may at least partially reflect increased attentional focus on nonlinguistic perceptual aspects of language. Hum Brain Mapp, 2013. © 2012 Wiley Periodicals, Inc.     

After-effects of 10 Hz tACS over the prefrontal cortex on phonological word decisions

IntroductionPrevious work in the language domain has shown that 10 Hz rTMS of the left or right posterior inferior frontal gyrus (pIFG) in the prefrontal cortex impaired phonological decision-making, arguing for a causal contribution of the bilateral pIFG to phonological processing. However, the neurophysiological correlates of these effects are unclear. The present study addressed the question whether neural activity in the prefrontal cortex could be modulated by 10 Hz tACS and how this would affect phonological decisions.MethodsIn three sessions, 24 healthy participants received tACS at 10 Hz or 16.18 Hz (control frequency) or sham stimulation over the bilateral prefrontal cortex before task processing. Resting state EEG was recorded before and after tACS. We also recorded EEG during task processing.ResultsRelative to sham stimulation, 10 Hz tACS significantly facilitated phonological response speed. This effect was task-specific as tACS did not affect a simple control task. Moreover, 10 Hz tACS significantly increased theta power during phonological decisions. The individual increase in theta power was positively correlated with the behavioral facilitation after 10 Hz tACS.ConclusionOur results show a facilitation of phonological decisions after 10 Hz tACS over the bilateral prefrontal cortex. This might indicate that 10 Hz tACS increased task-related activity in the stimulated area to a level that was optimal for phonological performance. The significant correlation with the individual increase in theta power suggests that the behavioral facilitation might be related to increased theta power during language processing.     

Functional links between motor and language systems

Transcranial magnetic stimulation (TMS) was applied to motor areas in the left language-dominant hemisphere while right-handed human subjects made lexical decisions on words related to actions. Response times to words referring to leg actions (e.g. kick) were compared with those to words referring to movements involving the arms and hands (e.g. pick). TMS of hand and leg areas influenced the processing of arm and leg words differentially, as documented by a significant interaction of the factors Stimulation site and Word category. Arm area TMS led to faster arm than leg word responses and the reverse effect, faster lexical decisions on leg than arm words, was present when TMS was applied to leg areas. TMS-related differences between word categories were not seen in control conditions, when TMS was applied to hand and leg areas in the right hemisphere and during sham stimulation. Our results show that the left hemispheric cortical systems for language and action are linked to each other in a category-specific manner and that activation in motor and premotor areas can influence the processing of specific kinds of words semantically related to arm or leg actions. By demonstrating specific functional links between action and language systems during lexical processing, these results call into question modular theories of language and motor functions and provide evidence that the two systems interact in the processing of meaningful information about language and action.     

Assembling and encoding word representations: fMRI subsequent memory effects implicate a role for phonological control

Novel word learning is central to the flexibility inherent in the human language capacity. Word learning may partially depend on long-term memory formation during the assembly of phonological representations from orthographic inputs. In the present study, event-related functional magnetic resonance imaging (fMRI) examined the contributions of phonological control-a component of the verbal working memory system-to phonological assembly and word learning. Subjects were scanned while making syllable decisions about visually presented familiar (English) and novel (pseudo-English and Foreign) words, a task that required retrieval and analysis of existing phonological codes or the assembly and analysis of novel representations. Results revealed that left inferior prefrontal cortex (LIPC) and bilateral parietal cortices were differentially engaged during the processing of novel words, suggesting that this circuit is recruited during phonological assembly. A subsequent memory analysis that examined the relation between fMRI signal and the subject's ability to later remember the words (a measure of effective memory formation) revealed that the magnitude of activation in LIPC, bilateral superior parietal, and left inferior parietal cortices was positively correlated with later memory. Moreover, although the magnitude of the subsequent memory effect in parietal cortex was not significantly affected by word type, this effect was greater in posterior LIPC for novel (pseudo-English) than for familiar (English) words. In the course of subserving the assembly of novel word representations, the phonological (articulatory) control component of the phonological system appears to play a central role in the encoding of novel words into long-term memory.     

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).     

Accessing orthographic representations from speech: The role of left ventral occipitotemporal cortex in spelling

The present fMRI study used a spelling task to investigate the hypothesis that the left ventral occipitotemporal cortex (vOT) hosts neuronal representations of whole written words. Such an orthographic word lexicon is posited by cognitive dual‐route theories of reading and spelling. In the scanner, participants performed a spelling task in which they had to indicate if a visually presented letter is present in the written form of an auditorily presented word. The main experimental manipulation distinguished between an orthographic word spelling condition in which correct spelling decisions had to be based on orthographic whole‐word representations, a word spelling condition in which reliance on orthographic whole‐word representations was optional and a phonological pseudoword spelling condition in which no reliance on such representations was possible. To evaluate spelling‐specific activations the spelling conditions were contrasted with control conditions that also presented auditory words and pseudowords, but participants had to indicate if a visually presented letter corresponded to the gender of the speaker. We identified a left vOT cluster activated for the critical orthographic word spelling condition relative to both the control condition and the phonological pseudoword spelling condition. Our results suggest that activation of left vOT during spelling can be attributed to the retrieval of orthographic whole‐word representations and, thus, support the position that the left vOT potentially represents the neuronal equivalent of the cognitive orthographic word lexicon. Hum Brain Mapp, 36:1393–1406, 2015. © 2014 The Authors Human Brain Mapping Published by Wiley Periodicals, Inc.     

Dissociating semantic and phonological contributions of the left inferior frontal gyrus to language production

While the involvement of the left inferior frontal gyrus (IFG) in language production is undisputed, the role of specific subregions at different representational levels remains unclear. Some studies suggest a division of anterior and posterior regions for semantic and phonological processing, respectively. Crucially, evidence thus far only comes from correlative neuroimaging studies, but the functional relevance of the involvement of these subregions during a given task remains elusive. We applied repetitive transcranial magnetic stimulation (rTMS) over anterior and posterior IFG (aIFG/pIFG), and vertex as a control site, while participants performed a category member and a rhyme generation task. We found a functional‐anatomical double dissociation between tasks and subregions. Naming latencies were significantly delayed in the semantic task when rTMS was applied to aIFG (relative to pIFG and vertex). In contrast, we observed a facilitation of naming latencies in the phonological task when rTMS was applied to pIFG (relative to aIFG and vertex). The results provide first causal evidence for the notion that anterior portions of the IFG are selectively recruited for semantic processing while posterior regions are functionally specific for phonological processing during word production. These findings shed light on the functional parcellation of the left IFG in language production.     

Reading in a regular orthography: an FMRI study investigating the role of visual familiarity

In order to separate the cognitive processes associated with phonological encoding and the use of a visual word form lexicon in reading, it is desirable to compare the processing of words presented in a visually familiar form with words in a visually unfamiliar form. Japanese Kana orthography offers this possibility. Two phonologically equivalent but visually dissimilar syllabaries allow the writing of, for example, foreign loanwords in two ways, only one of which is visually familiar. Familiarly written words, unfamiliarly written words, and pseudowords were presented in both Kana syllabaries (yielding six conditions in total) to participants during an fMRI measurement with a silent articulation task (Experiment 1) and a phonological lexical decision task (Experiment 2) using an event-related design. Consistent over two experimental tasks, the three different stimulus types (familiar, unfamiliar, and pseudoword) were found to activate selectively different brain regions previously associated with phonological encoding and word retrieval or meaning. Compatible with the predictions of the dual-route model for reading, pseudowords and visually unfamiliar words, which have to be read using phonological assembly, caused an increase in brain activity in left inferior frontal regions (BA 44/47), as compared to visually familiar words. Visually familiar and unfamiliar words were found to activate a range of areas associated with lexico-semantic processing more strongly than pseudowords, such as the left and right temporo-parietal region (BA 39/40), a region in the left middle/inferior temporal gyrus (BA 20/21), and the posterior cingulate (BA 31).     

Single‐word semantic judgements in semantic dementia: Do phonology and grammatical class count?

Background: Listeners make active use of phonological regularities such as word length to facilitate higher‐level syntactic and semantic processing. For example, nouns are longer than verbs, and abstract words are longer than concrete words. Patients with semantic dementia (SD) experience conceptual loss with preserved syntax and phonology. The extent to which patients with SD exploit phonological regularities to support language processing remains unclear.

Aims: We examined the ability of patients with SD (1) to perceive subtle acoustic–phonetic distinctions in English, and (2) to bootstrap their accuracy of lexical‐semantic and syntactic judgements from regularities in the phonological forms of English nouns and verbs.

Methods and Procedures: Four patients with SD made minimal pair judgements (same/different) for auditorily presented stimuli selectively varied by voice, place, or manner of the initial consonant (e.g., pa –ba). In Experiment 2 patients made forced‐choice semantic judgements (abstract or concrete) for single words varied by (1) concreteness (abstract or concrete); (2) grammatical class (noun or verb); and (3) word length (one‐ or three‐syllable words).

Outcomes and Results: The most semantically impaired patients paradoxically showed the highest accuracy of minimal pair phonologic discrimination. Judgements of word concreteness were less accurate for verbs than nouns. Among verbs, accuracy was worse for concrete than abstract items (e.g., eat was worse than think). Patients were more likely to misclassify longer concrete words (e.g., professor) as abstract, demonstrating sensitivity to an underlying phonologically mediated semantic property in English.

Conclusions: Single‐word semantic judgements were sensitive to both grammatical class and phonological properties of the words being evaluated. Theoretical and clinical implications are addressed in the context of an anatomically constrained model of SD that assumes increasing reliance on phonology as lexical‐semantic knowledge degrades.     

Brain regions associated with successful and unsuccessful retrieval of verbal episodic memory as revealed by divided attention

Which brain regions are implicated when words are retrieved under divided attention, and what does this tell us about attentional and memory processes needed for retrieval? To address these questions we used fMRI to examine brain regions associated with auditory recognition performed under full and divided attention (DA). We asked young adults to encode words presented auditorily under full attention (FA), and following this, asked them to recognize studied words while in the scanner. Attention was divided at retrieval by asking participants to perform either an animacy task to words, or odd-digit identification task to numbers presented visually, concurrently with the recognition task. Retrieval was disrupted significantly by the word-, but not number-based concurrent task. A corresponding decrease in brain activity was observed in right hippocampus, bilateral parietal cortex, and left precuneus, thus demonstrating, for the first time, involvement of these regions in recognition under DA at retrieval. Increases in activation of left prefrontal cortex (PFC), associated with phonological processing, were observed in the word- compared to number-based DA condition. Results suggest that the medial temporal lobe (MTL) and neo-cortical components of retrieval, believed to form the basis of episodic memory traces, are disrupted when phonological processing regions in left PFC are engaged simultaneously by another task. Results also support a component-process model of retrieval which posits that MTL-mediated retrieval does not compete for general cognitive resources but does compete for specific structural representations.     

Reliability of the 'observation of movement' method for determining motor threshold using transcranial magnetic stimulation

The aim of this study was to establish the reliability of the observation of movement (OM) method for obtaining motor threshold (MT) in transcranial magnetic stimulation (TMS). MTs were obtained on separate days, following separate hunting procedures, for both left and right motor cortex (M1), with one or multiple estimates obtained from the same hemisphere within a single session. MTs obtained using the OM method were highly reliable and reproducible on different days (left M1: r = .98, p < .0001; right M1: r = .97, p < .0001). MTs were not influenced by the order of acquisition when two hemispheres were stimulated in the same session [F(1,22) = .12, p = .73] or by the collection of additional MTs as part of the distance-adjusted procedure [F(1,23) = .74, p = .40]. The results verify the reliability of the OM method and confirm its viability for the safe and efficient application of TMS to the left and right M1. The OM method is a reliable technique for obtaining MT and is relatively simple and quick to run. It therefore provides an effective procedure for research and clinical applications.     

Cortical activation during word processing in late bilinguals: similarities and differences as revealed by functional magnetic resonance imaging

Functional magnetic resonance imaging was used to compare cortical organization of the first (L1, Russian) and second (L2, English) languages. Six fluent Russian-English bilinguals who acquired their second language postpuberty were tested with words and nonwords presented either auditorily or visually. Results showed that both languages activated similar cortical networks, including the inferior frontal, middle frontal, superior temporal, middle temporal, angular, and supramarginal gyri. Within the inferior frontal gyrus (IFG), L2 activated a larger cortical volume than L1 during lexical and phonological processing. For both languages, the left IFG was more active than the right IFG during lexical processing. Within the left IFG, the distance between centers of activation associated with lexical processing of translation equivalents across languages was larger than the distance between centers of activation associated with lexical processing of different words in the same language. Results of phonological processing analyses revealed different centers of activation associated with the first versus the second language in the IFG, but not in the superior temporal gyrus (STG). These findings are discussed within the context of the current literature on cortical organization in bilinguals and suggest variation in bilingual cortical activation associated with lexical, phonological, and orthographic processing.     

Lexical ambiguity resolution in Wernicke's area and its right homologue

Introduction

There is an academic dispute regarding the role of the right hemisphere in language processing. Transcranial Magnetic Stimulation (TMS) was used to test the hypothesis that Wernicke's area processes dominant meanings (“teller”) whereas its right homologue processes subordinate meanings (“river”) of ambiguous words (“bank”; Jung-Beeman, 2005).

Methods

Participants were asked to make a semantic decision on ambiguous words that were followed either by unrelated words or by words associated with their dominant or subordinate meanings. A 10 Hz TMS train was applied on each trial over CP5 (left Wernicke), CP6 (right Wernicke) or Cz (vertex) scalp positions, and was synchronized with the word presentation.

Results

Accuracy and d′ analysis revealed a TMS LOCATION by MEANING interaction. TMS over Wernicke's area resulted in more accurate responses and higher sensitivity to dominant meaning blocks compared to stimulating the right Wernicke's area and the vertex. In contrast, TMS over the right Wernicke's area resulted in more accurate responses and higher sensitivity to subordinate meaning blocks, compared to stimulating the left Wernicke's area and the vertex.

Conclusion

The left and right Wernicke's areas function as processors of dominant and subordinate meanings of ambiguous words, respectively. While previous research methods have yielded indecisive results, TMS proved to be a useful tool in demonstrating a causal role of the two brain regions in a double dissociation design with healthy subjects.     

Functional brain network abnormalities during verbal working memory performance in adolescents and young adults with dyslexia

Behavioral and functional neuroimaging studies indicate deficits in verbal working memory (WM) and frontoparietal dysfunction in individuals with dyslexia. Additionally, structural brain abnormalities in dyslexics suggest a dysconnectivity of brain regions associated with phonological processing. However, little is known about the functional neuroanatomy underlying cognitive dysfunction in dyslexia. In this study, functional magnetic resonance imaging and multivariate analytic techniques were used to investigate patterns of functional connectivity during a verbal WM task in individuals with dyslexia (n = 12) and control subjects (n = 13). Dyslexics were not significantly slower than controls; however, they were less accurate with increasing WM demand. Independent component analysis identified 18 independent components (ICs) among which two ICs were selected for further analyses. These ICs included functional networks which were positively correlated with the delay period of the activation task in both healthy controls and dyslexics. Connectivity abnormalities in dyslexics were detected within both networks of interest: within a “phonological” left-lateralized prefrontal network, increased functional connectivity was found in left prefrontal and inferior parietal regions. Within an “executive” bilateral frontoparietal network, dyslexics showed a decreased connectivity pattern comprising bilateral dorsolateral prefrontal and posterior parietal regions, while increased connectivity was found in the left angular gyrus, the left hippocampal cortex and the right thalamus. The functional connectivity strength in the latter regions was associated with WM task accuracy and with the numbers of errors during a spelling test. These data suggest functional connectivity abnormalities in two spatiotemporally dissociable brain networks underlying WM dysfunction in individuals with dyslexia.     

The effect of phonological repetition on cortical magnetic responses evoked by visually presented words

Neuroimaging studies have reported that the left superior temporal cortical area is activated by visually presented words. In the present study, we recorded cortical magnetic responses evoked by visual words and examined the effect of phonological repetition (e.g., hair-hare) on left superior temporal cortical activity, using pairs of homophonic Japanese words as stimuli. Unlike English, Japanese has a large number of homophone pairs with a totally different orthography. By taking advantage of this feature of the Japanese writing system, the effect of phonological repetition can be solely examined without being confounded by the effect of orthographic similarity. Magnetic responses were recorded over the bilateral temporal sites of the brain while subjects silently read words. The words were presented one by one; a quarter of them was immediately followed by a homophonic word. Clear magnetic responses in the latency range of 300-600 msec were observed in the left hemisphere, and the responses to the homophones were smaller than those to the first presented words. In the right hemisphere, clear responses were not consistently recorded in the same latency range, and no effect of phonological repetition was observed. The sources of the responses recorded over the left hemisphere were estimated to be in the left superior temporal cortical area adjacent to the auditory cortex and the source strength as well as the magnetic responses showed a reduction by phonological repetition. This result suggests that the activity in the left superior temporal cortical area is associated with access to the phonological representation of words.     

Cortical activation patterns during long-term memory retrieval of visually or haptically encoded objects and locations

The present study used functional magnetic resonance imaging to delineate cortical networks that are activated when objects or spatial locations encoded either visually (visual encoding group, n=10) or haptically (haptic encoding group, n=10) had to be retrieved from long-term memory. Participants learned associations between auditorily presented words and either meaningless objects or locations in a 3-D space. During the retrieval phase one day later, participants had to decide whether two auditorily presented words shared an association with a common object or location. Thus, perceptual stimulation during retrieval was always equivalent, whereas either visually or haptically encoded object or location associations had to be reactivated. Moreover, the number of associations fanning out from each word varied systematically, enabling a parametric increase of the number of reactivated representations. Recall of visual objects predominantly activated the left superior frontal gyrus and the intraparietal cortex, whereas visually learned locations activated the superior parietal cortex of both hemispheres. Retrieval of haptically encoded material activated the left medial frontal gyrus and the intraparietal cortex in the object condition, and the bilateral superior parietal cortex in the location condition. A direct test for modality-specific effects showed that visually encoded material activated more vision-related areas (BA 18/19) and haptically encoded material more motor and somatosensory-related areas. A conjunction analysis identified supramodal and material-unspecific activations within the medial and superior frontal gyrus and the superior parietal lobe including the intraparietal sulcus. These activation patterns strongly support the idea that code-specific representations are consolidated and reactivated within anatomically distributed cell assemblies that comprise sensory and motor processing systems.     

Neural pathways involved in the processing of concrete and abstract words.

The purpose of this study was to delineate the neural pathways involved in processing concrete and abstract words using functional magnetic resonance imaging (fMRI). Word and pseudoword stimuli were presented visually, one at a time, and the participant was required to make a lexical decision. Lexical decision epochs alternated with a resting baseline. In each lexical decision epoch, the stimuli were either concrete words and pseudowords, or abstract words and pseudowords. Behavioral data indicated that, as with previous research, concrete word stimuli were processed more efficiently than abstract word stimuli. Analysis of the fMRI data indicated that processing of word stimuli, compared to the baseline condition, was associated with neural activation in the bilateral fusiform gyrus, anterior cingulate, left middle temporal gyrus, right posterior superior temporal gyrus, and left and right inferior frontal gyrus. A direct comparison between the abstract and concrete stimuli epochs yielded a significant area of activation in the right anterior temporal cortex. The results are consistent with recent positron emission tomography work showing right hemisphere activation during processing of abstract representations of language. The results are interpreted as support for a right hemisphere neural pathway in the processing of abstract word representations.     

fMRI evidence for dual routes to the mental lexicon in visual word recognition.

Event-related fMRI was used to investigate lexical decisions to words of high and low frequency of occurrence and to pseudowords. The results obtained strongly support dual-route models of visual word processing. By contrasting words with pseudowords, bilateral occipito-temporal brain areas and posterior left middle temporal gyrus (MTG) were identified as contributing to the successful mapping of orthographic percepts onto visual word form representations. Low-frequency words and pseudowords elicited greater activations than high-frequency words in the superior pars opercularis [Brodmann's area (BA) 44] of the left inferior frontal gyrus (IFG), in the anterior insula, and in the thalamus and caudate nucleus. As processing of these stimuli during lexical search is known to rely on phonological information, it is concluded that these brain regions are involved in grapheme-to-phoneme conversion. Activation in the pars triangularis (BA 45) of the left IFG was observed only for low-frequency words. It is proposed that this region is involved in processes of lexical selection.