Left Ventral Occipitotemporal Cortex Research Articles

Orthographic and phonological processing in the left ventral occipitotemporal cortex during Chinese word reading.

The left ventral occipitotemporal cortex (lvOT) has been consistently identified as a crucial structure in word reading, and its function varies across subregions. Nevertheless, the specific function of the lvOT and its subregions remains controversial because the obvious grapheme-to-phoneme correspondence rules of alphabetic languages make it difficult to disentangle the contributions of orthography and phonology to neural activations. To explore information processing in lvOT subregions, the present study manipulated the orthography and phonology in a factorial design and used the fMRI rapid adaptation paradigm. The results revealed a posterior-to-anterior functional gradient in lvOT in Chinese word reading and specified that the functional transition from sublexical to lexical processing occurred in the middle subregion close to the classic VWFA. More importantly, we found that the middle and posterior subregions of lvOT are responsible for processing both orthographic and phonological information during Chinese word reading. These results elaborated the function of the lvOT in Chinese word reading.

Reading Reshapes Stimulus Selectivity in the Visual Word Form Area.

Reading depends on a brain region known as the "visual word form area" (VWFA) in the left ventral occipitotemporal cortex. This region's function is debated because its stimulus selectivity is not absolute, it is modulated by a variety of task demands, and it is inconsistently localized. We used fMRI to characterize the combination of sensory and cognitive factors that activate word-responsive regions that we precisely localized in 16 adult humans (4 male). We then presented three types of character strings: English words, pseudowords, and unfamiliar characters with matched visual features. Participants performed three different tasks while viewing those stimuli: detecting real words, detecting color in the characters, and detecting color in the fixation mark. There were three primary findings about the VWFA's response: (1) It preferred letter strings over unfamiliar characters even when the stimuli were ignored during the fixation task. (2) Compared with those baseline responses, engaging in the word reading task enhanced the response to words but suppressed the response to unfamiliar characters. (3) Attending to the stimuli to judge their color had little effect on the response magnitudes. Thus, the VWFA is uniquely modulated by a cognitive signal that is specific to voluntary linguistic processing and is not additive. Functional connectivity analyses revealed that communication between the VWFA and a left frontal language area increased when the participant engaged in the linguistic task. We conclude that the VWFA is inherently selective for familiar orthography, but it falls under control of the language network when the task demands it.

Reading reshapes stimulus selectivity in the visual word form area.

Reading depends on a brain region known as the "visual word form area" (VWFA) in left ventral occipito-temporal cortex. This region's function is debated because its stimulus selectivity is not absolute, it is modulated by a variety of task demands, and it is inconsistently localized. We used fMRI to characterize the combination of sensory and cognitive factors that activate word-responsive regions that we precisely localized in 16 adult humans (4 male). We then presented three types of character strings: English words, pseudowords, and unfamiliar characters with matched visual features. Participants performed three different tasks while viewing those stimuli: detecting real words, detecting color in the characters, and detecting color in the fixation mark. There were three primary findings about the VWFA's response: (1) It preferred letter strings over unfamiliar characters even when the stimuli were ignored during the fixation task; (2) Compared to those baseline responses, engaging in the word reading task enhanced the response to words but suppressed the response to unfamiliar characters. (3) Attending to the stimuli to judge their font color had little effect on the response magnitudes. Thus, the VWFA is uniquely modulated by a cognitive signal that is specific to voluntary linguistic processing and is not additive. Functional connectivity analyses revealed that communication between the VWFA and a left frontal language area increased when the participant engaged in the linguistic task. We conclude that the VWFA is inherently selective for familiar orthography, but it falls under control of the language network when the task demands it.

The role of the left ventral occipitotemporal cortex in speech processing-The influence of visual deprivation.

The role of the left ventral occipitotemporal cortex (vOT) in reading is well-established in both sighted and blind readers. Its role in speech processing remains only partially understood. Here, we test the involvement of the left vOT in phonological processing of spoken language in the blind (N = 50, age: 6.76-60.32) and in the sighted (N = 54, age: 6.79-59.83) by means of whole-brain and region-of-interest (including individually identified) fMRI analyses. We confirm that the left vOT is sensitive to phonological processing (shows greater involvement in rhyming compared to control spoken language task) in both blind and sighted participants. However, in the sighted, the activation was observed only during the rhyming task and in the speech-specific region of the left vOT, pointing to task and modality specificity. In contrast, in the blind group, the left vOT was active during speech processing irrespective of task and in both speech and reading-specific vOT regions. Only in the blind, the left vOT presented a higher degree of sensitivity to phonological processing than other language nodes in the left inferior frontal and superior temporal cortex. Our results suggest a changed development of the left vOT sensitivity to spoken language, resulting from visual deprivation.

Adaptive short-term plasticity in the typical reading network

The left temporo-parietal cortex (TPC) is crucial for phonological decoding, i.e., for learning and retaining sound-letter mappings, and appears hypoactive in dyslexia. Here, we tested the causal contribution of this area for reading in typical readers with transcranial magnetic stimulation (TMS) and explored the reading network's response with fMRI. By investigating the underlying neural correlates of stimulation-induced modulations of the reading network, we can help improve targeted interventions for individuals with dyslexia. 28 typical adult readers overtly read simple and complex words and pseudowords during fMRI after effective and sham TMS over the left TPC. To explore differences in functional activation and effective connectivity within the reading network, we performed univariate and multivariate analyses, as well as dynamic causal modeling. While TMS-induced effects on reading performance and brain activation showed large individual variability, multivariate analyses revealed a shift in activation in the left inferior frontal cortex for pseudoword reading after effective TMS. Furthermore, TMS increased effective connectivity from the left ventral occipito-temporal cortex to the left TPC. In the absence of effects on reading performance, the observed changes in task-related activity and the increase in functional coupling between the two core reading nodes suggest successful short-term compensatory reorganization in the reading network following TMS-induced disruption. This study is the first to explore neurophysiological changes induced by TMS to a core reading node in typical readers while performing an overt reading task. We provide evidence for remote stimulation effects and emphasize the relevance of functional interactions in the reading network.

Beyond the Visual Word Form Area - a cognitive characterization of the left ventral occipitotemporal cortex.

The left ventral occipitotemporal cortex has been traditionally viewed as a pathway for visual object recognition including written letters and words. Its crucial role in reading was strengthened by the studies on the functionally localized "Visual Word Form Area" responsible for processing word-like information. However, in the past 20 years, empirical studies have challenged the assumptions of this brain region as processing exclusively visual or even orthographic stimuli. In this review, we aimed to present the development of understanding of the left ventral occipitotemporal cortex from the visually based letter area to the modality-independent symbolic language related region. We discuss theoretical and empirical research that includes orthographic, phonological, and semantic properties of language. Existing results showed that involvement of the left ventral occipitotemporal cortex is not limited to unimodal activity but also includes multimodal processes. The idea of the integrative nature of this region is supported by the broad functional and structural connectivity with language-related and attentional brain networks. We conclude that although the function of the area is not yet fully understood in human cognition, its role goes beyond visual word form processing. The left ventral occipitotemporal cortex seems to be crucial for combining higher-level language information with abstract forms that convey meaning independently of modality.

Visual event-related potentials reveal the early whole-word lexical processing of Chinese two-character words

Morphologically complex words are common across different languages, especially in Chinese, because more than 90% of common modern Chinese words are complex words. Many behavioral studies have suggested the whole-word processing of Chinese complex words, but the neural correlates of whole-word processing remain unclear. Previous electrophysiological studies revealed automatic and early (∼250 ms) access to the orthographic forms of monomorphic words in the ventral occipitotemporal area. In this study, we investigated whether there is also automatic and early orthographic recognition of Chinese complex words (as whole units) by recording event-related potentials (ERPs). A total of 150 two-character words and 150 two-character pseudowords composed of the same 300 characters (morphemes) were pseudorandomly presented to proficient Chinese readers. Participants were required to determine the color of each stimulus in the color decision task and to determine whether each stimulus was a word in the lexical decision task. The two constituent characters of each stimulus were horizontally arranged in Experiment 1 and vertically arranged in Experiment 2. The results revealed a significant early ERP difference between words and pseudowords approximately 250–300 ms after stimulus onset in the parieto-occipital scalp region. The early ERP difference was more prominent in the color decision task than in the lexical decision task, more prominent in Experiment 1 than in Experiment 2, and more prominent in the left parieto-occipital scalp region than in the right. Source analysis results showed that the early ERP difference originated from the left ventral occipitotemporal cortex. These results reflected early and automatic access to whole-word orthographic representations of Chinese complex words in the left ventral occipitotemporal cortex.

Neural attenuation: age-related dedifferentiation in the left occipitotemporal cortex for visual word processing.

The present study investigated the age-related neural basis of cognitive decline in the left ventral occipitotemporal cortex (vOT)-a brain area that responds selectively to visual words processing. Functional magnetic resonance imaging was used to estimate neural activity in this area, while young and old adults viewed words and line drawings. Our results demonstrated the existence of neural dedifferentiation of the left vOT in old adults during visual word processing. More specifically, this dedifferentiation was due to neural attenuation that is, decreased response to words rather than increased response to line drawings in old adults compared with young adults. In addition, individuals who showed decreased neural response to words had worse performance in visual word processing. Taken together, our findings support the neural attenuation hypothesis for the cognitive decline in visual word processing in old adults.

Graph theoretical analysis reveals the functional role of the left ventral occipito-temporal cortex in speech processing

The left ventral occipito-temporal cortex (left-vOT) plays a key role in reading. Interestingly, the area also responds to speech input, suggesting that it may have other functions beyond written word recognition. Here, we adopt graph theoretical analysis to investigate the left-vOT’s functional role in the whole-brain network while participants process spoken sentences in different contexts. Overall, different connectivity measures indicate that the left-vOT acts as an interface enabling the communication between distributed brain regions and sub-networks. During simple speech perception, the left-vOT is systematically part of the visual network and contributes to the communication between neighboring areas, remote areas, and sub-networks, by acting as a local bridge, a global bridge, and a connector, respectively. However, when speech comprehension is explicitly required, the specific functional role of the area and the sub-network to which the left-vOT belongs change and vary with the quality of speech signal and task difficulty. These connectivity patterns provide insightful information on the contribution of the left-vOT in various contexts of language processing beyond its role in reading. They advance our general understanding of the neural mechanisms underlying the flexibility of the language network that adjusts itself according to the processing context.

Effective connectivity of the left-ventral occipito-temporal cortex during visual word processing: Direct causal evidence from TMS-EEG co-registration

As an interface between the visual and language system, the left ventral occipito-temporal cortex (left-vOT) plays a key role in reading. This functional role is supported by anatomical and functional connections between the area and other brain regions within and outside the language network. Nevertheless, only a few studies have investigated how the functional state of this area, which is dependent upon the nature of the task demand and the stimulus being processed, could influence the activity of the connected brain regions. In the present combined TMS-EEG study, we studied the left-vOT effective connectivity by adopting a direct, causal intervention approach. Using TMS, we probed left-vOT activation in different processing contexts and measured the neural propagation of activity from this area to other brain regions. A comparison of neural propagation measured during low-level visual detection of language versus non-language stimuli showed that processing language stimuli reduced neural propagation from the left-vOT to the right occipital cortex. Additionally, compared to the low-level visual detection of language stimuli, performing semantic judgments on the same stimuli further reduced neural propagation to the posterior part of the corpus callosum, right superior parietal lobule and the right anterior temporal lobe. This reduction of cross-hemispheric neural propagation was accompanied by an increase in the collaboration between areas within the left-hemisphere language network. Together, this first evidence from a direct causal intervention approach suggests that processing language stimuli and performing a high-level language task reduce effective connectivity from the left-vOT to the right hemisphere, and may contribute to the left-hemisphere lateralization typically observed during language processing.

Task modulates the orthographic and phonological representations in the bilateral ventral Occipitotemporal cortex.

As a key area in word reading, the left ventral occipitotemporal cortex is proposed for abstract orthographic processing, and its middle part has even been labeled as the visual word form area. Because the definition of the VWFA largely varies and the reading task differs across studies, the function of the left ventral occipitotemporal cortex in word reading is continuingly debated on whether this region is specific for orthographic processing or be involved in an interactive framework. By using representational similarity analysis (RSA), this study examined information representation in the VWFA at the individual level and the modulatory effect of reading task. Twenty-four subjects were scanned while performing the explicit (i.e., the naming task) and implicit (i.e., the perceptual task) reading tasks. Activation analysis showed that the naming task elicited greater activation in regions related to phonological processing (e.g., the bilateral prefrontal cortex and temporoparietal cortex), while the perceptual task recruited greater activation in visual cortex and default mode network (e.g., the bilateral middle frontal gyrus, angular gyrus, and the right middle temporal gyrus). More importantly, RSA also showed that task modulated information representation in the bilateral anterior occipitotemporal cortex and VWFA. Specifically, ROI-based RSA revealed enhanced orthographic and phonological representations in the bilateral anterior fusiform cortex and VWFA in the naming task relative to the perceptual task. These results suggest that lexical representation in the VWFA is influenced by the demand of phonological processing, which supports the interactive account of the VWFA.

Task-Relevant Representations and Cognitive Control Demands Modulate Functional Connectivity from Ventral Occipito-Temporal Cortex During Object Recognition Tasks.

The left ventral occipito-temporal cortex (vOTC) supports extraction and processing of visual features. However, it has remained unclear whether left vOTC-based functional connectivity (FC) differs according to task-relevant representations (e.g., lexical, visual) and control demands imposed by the task, even when similar visual-semantic processing is required for object identification. Here, neural responses to the same set of pictures of meaningful objects were measured, while the type of task that participants had to perform (picture naming versus size-judgment task), and the level of cognitive control required by the picture naming task (high versus low interference contexts) were manipulated. Explicit retrieval of lexical representations in the picture naming task facilitated activation of lexical/phonological representations, modulating FC between left vOTC and dorsal anterior-cingulate-cortex/pre-supplementary-motor-area. This effect was not observed in the size-judgment task, which did not require explicit word-retrieval of object names. Furthermore, retrieving the very same lexical/phonological representation in the high versus low interference contexts during picture naming increased FC between left vOTC and left caudate. These findings support the proposal that vOTC functional specialization emerges from interactions with task-relevant brain regions.

Is human face recognition lateralized to the right hemisphere due to neural competition with left-lateralized visual word recognition? A critical review.

The right hemispheric lateralization of face recognition, which is well documented and appears to be specific to the human species, remains a scientific mystery. According to a long-standing view, the evolution of language, which is typically substantiated in the left hemisphere, competes with the cortical space in that hemisphere available for visuospatial processes, including face recognition. Over the last decade, a specific hypothesis derived from this view according to which neural competition in the left ventral occipito-temporal cortex with selective representations of letter strings causes right hemispheric lateralization of face recognition, has generated considerable interest and research in the scientific community. Here, a systematic review of studies performed in various populations (infants, children, literate and illiterate adults, left-handed adults) and methodologies (behavior, lesion studies, (intra)electroencephalography, neuroimaging) offers little if any support for this reading lateralized neural competition hypothesis. Specifically, right-lateralized face-selective neural activity already emerges at a few months of age, well before reading acquisition. Moreover, consistent evidence of face recognition performance and its right hemispheric lateralization being modulated by literacy level during development or at adulthood is lacking. Given the absence of solid alternative hypotheses and the key role of neural competition in the sensory-motor cortices for selectivity of representations, learning, and plasticity, a revised language-related neural competition hypothesis for the right hemispheric lateralization of face recognition should be further explored in future research, albeit with substantial conceptual clarification and advances in methodological rigor.

Distinct neural sources underlying visual word form processing as revealed by steady state visual evoked potentials (SSVEP)

EEG has been central to investigations of the time course of various neural functions underpinning visual word recognition. Recently the steady-state visual evoked potential (SSVEP) paradigm has been increasingly adopted for word recognition studies due to its high signal-to-noise ratio. Such studies, however, have been typically framed around a single source in the left ventral occipitotemporal cortex (vOT). Here, we combine SSVEP recorded from 16 adult native English speakers with a data-driven spatial filtering approach—Reliable Components Analysis (RCA)—to elucidate distinct functional sources with overlapping yet separable time courses and topographies that emerge when contrasting words with pseudofont visual controls. The first component topography was maximal over left vOT regions with a shorter latency (approximately 180 ms). A second component was maximal over more dorsal parietal regions with a longer latency (approximately 260 ms). Both components consistently emerged across a range of parameter manipulations including changes in the spatial overlap between successive stimuli, and changes in both base and deviation frequency. We then contrasted word-in-nonword and word-in-pseudoword to test the hierarchical processing mechanisms underlying visual word recognition. Results suggest that these hierarchical contrasts fail to evoke a unitary component that might be reasonably associated with lexical access.

Neural network for Braille reading and the speech-reading convergence in the blind: Similarities and differences to visual reading

All writing systems represent units of spoken language. Studies on the neural correlates of reading in different languages show that this skill relies on access to brain areas dedicated to speech processing. Speech-reading convergence onto a common perisylvian network is therefore considered universal among different writing systems. Using fMRI, we test whether this holds true also for tactile Braille reading in the blind. The neural networks for Braille and visual reading overlapped in the left ventral occipitotemporal (vOT) cortex. Even though we showed similar perisylvian specialization for speech in both groups, blind subjects did not engage this speech system for reading. In contrast to the sighted, speech-reading convergence in the blind was absent in the perisylvian network. Instead, the blind engaged vOT not only in reading but also in speech processing. The involvement of the vOT in speech processing and its engagement in reading in the blind suggests that vOT is included in a modality independent language network in the blind, also evidenced by functional connectivity results. The analysis of individual speech-reading convergence suggests that there may be segregated neuronal populations in the vOT for speech processing and reading in the blind.

Chapter 12 - Reading and alexia

Alexia refers to a reading disorder caused by some form of acquired brain pathology, most commonly a stroke or tumor, in a previously literate subject. In neuropsychology, a distinction is made between central alexia (commonly seen in aphasia) and peripheral alexia (a perceptual or attentional deficit). The prototypical peripheral alexia is alexia without agraphia (pure alexia), where patients can write but are impaired in reading words and letters. Pure alexia is associated with damage to the left ventral occipitotemporal cortex (vOT) or its connections. Hemianopic alexia is associated with less extensive occipital damage and is caused by a visual field defect, which creates problems reading longer words and passages of text. Reading impairment can also arise due to attentional deficits, most commonly following right hemisphere or bilateral lesions. Studying patients with alexia, along with functional imaging studies of normal readers, has improved our understanding of the neurobiological processes involved in reading. A key question is whether an area in the left ventral occipitotemporal cortex is specialized for or selectively involved in word processing, or whether reading relies on tuning of more general purpose perceptual areas. Reading deficits may also be observed in dementia and traumatic brain injury, but often with less consistent deficit patterns than in patients with focal lesions.

Letter fluency in 7-8-year-old children is related to the anterior, but not posterior, ventral occipito-temporal cortex during an auditory phonological task

Previous studies have shown that reading skill in 3- to 6-year-old children is related to the automatic activation of the posterior left ventral occipitotemporal cortex (vOT) during spoken language processing, whereas 8- to 15-year-old children and adult readers activate the anterior vOT. However, it is unknown how children who are between these two age groups automatically activate orthographic representations in vOT for spoken language. In the current study, we recruited 153 7- to 8-year-old children to fill the age gap from previous studies. Using functional magnetic resonance imaging (fMRI), we measured children’s reading-related skills and brain activity during an auditory phonological task with both a small (i.e. onset) and a large (i.e. rhyme) grain size condition. We found that letter fluency, but not reading accuracy, was correlated with activation in the anterior vOT for the rhyme condition. There were no reading-related skill correlations for the posterior vOT or for activation during the onset condition in this age group. Our findings reveal that automatic activation in the anterior vOT during spoken language processing already occurs in higher skilled 7- to 8-year-old children. In addition, increases in naming automaticity is the primary determinant of the engagement of vOT during phonological awareness tasks.

Spatiotemporal dynamics of orthographic and lexical processing in the ventral visual pathway.

Reading is a rapid, distributed process that engages multiple components of the ventral visual stream. To understand the neural constituents and their interactions that allow us to identify written words, we performed direct intra-cranial recordings in a large cohort of humans. This allowed us to isolate the spatiotemporal dynamics of visual word recognition across the entire left ventral occipitotemporal cortex. We found that mid-fusiform cortex is the first brain region sensitive to lexicality, preceding the traditional visual word form area. The magnitude and duration of its activation are driven by the statistics of natural language. Information regarding lexicality and word frequency propagates posteriorly from this region to visual word form regions and to earlier visual cortex, which, while active earlier, show sensitivity to words later. Further, direct electrical stimulation of this region results in reading arrest, further illustrating its crucial role in reading. This unique sensitivity of mid-fusiform cortex to sub-lexical and lexical characteristics points to its central role as the orthographic lexicon-the long-term memory representations of visual word forms.

Visual word form processing deficits driven by severity of reading impairments in children with developmental dyslexia

The visual word form area (VWFA) in the left ventral occipito-temporal (vOT) cortex is key to fluent reading in children and adults. Diminished VWFA activation during print processing tasks is a common finding in subjects with severe reading problems. Here, we report fMRI data from a multicentre study with 140 children in primary school (7.9–12.2 years; 55 children with dyslexia, 73 typical readers, 12 intermediate readers). All performed a semantic task on visually presented words and a matched control task on symbol strings. With this large group of children, including the entire spectrum from severely impaired to highly fluent readers, we aimed to clarify the association of reading fluency and left vOT activation during visual word processing. The results of this study confirm reduced word-sensitive activation within the left vOT in children with dyslexia. Interestingly, the association of reading skills and left vOT activation was especially strong and spatially extended in children with dyslexia. Thus, deficits in basic visual word form processing increase with the severity of reading disability but seem only weakly associated with fluency within the typical reading range suggesting a linear dependence of reading scores with VFWA activation only in the poorest readers.

Differential activation of the visual word form area during auditory phoneme perception in youth with dyslexia

Developmental dyslexia is a learning disorder characterized by difficulties reading words accurately and/or fluently. Several behavioral studies have suggested the presence of anomalies at an early stage of phoneme processing, when the complex spectrotemporal patterns in the speech signal are analyzed and assigned to phonemic categories. In this study, fMRI was used to compare brain responses associated with categorical discrimination of speech syllables (P) and acoustically matched nonphonemic stimuli (N) in children and adolescents with dyslexia and in typically developing (TD) controls, aged 8–17 years. The TD group showed significantly greater activation during the P condition relative to N in an area of the left ventral occipitotemporal cortex that corresponds well with the region referred to as the “visual word form area” (VWFA). Regression analyses using reading performance as a continuous variable across the full group of participants yielded similar results. Overall, the findings are consistent with those of previous neuroimaging studies using print stimuli in individuals with dyslexia that found reduced activation in left occipitotemporal regions; however, the current study shows that these activation differences seen during reading are apparent during auditory phoneme discrimination in youth with dyslexia, suggesting that the primary deficit in at least a subset of children may lie early in the speech processing stream and that categorical perception may be an important target of early intervention in children at risk for dyslexia.