A dynamic causal modeling analysis of the effective connectivities underlying top-down letter processing

The present study employed dynamic causal modeling to investigate the effective functional connectivity between regions of the neural network involved in top-down letter processing. We used an illusory letter detection paradigm in which participants detected letters while viewing pure noise images. When participants detected letters, the response of the right middle occipital gyrus (MOG) in the visual cortex was enhanced by increased feed-backward connectivity from the left inferior frontal gyrus (IFG). In addition, illusory letter detection increased feed-forward connectivity from the right MOG to the left inferior parietal lobules. Originating in the left IFG, this top-down letter processing network may facilitate the detection of letters by activating letter processing areas within the visual cortex. This activation in turns may highlight the visual features of letters and send letter information to activate the associated phonological representations in the identified parietal region.     

Neural correlates of top-down letter processing

This fMRI study investigated top-down letter processing with an illusory letter detection task. Participants responded whether one of a number of different possible letters was present in a very noisy image. After initial training that became increasingly difficult, they continued to detect letters even though the images consisted of pure noise, which eliminated contamination from strong bottom-up input. For illusory letter detection, greater fMRI activation was observed in several cortical regions. These regions included the precuneus, an area generally involved in top-down processing of objects, and the left superior parietal lobule, an area previously identified with the processing of valid letter and word stimuli. In addition, top-down letter detection also activated the left inferior frontal gyrus, an area that may be involved in the integration of general top-down processing and letter-specific bottom-up processing. These findings suggest that these regions may play a significant role in top-down as well as bottom-up processing of letters and words, and are likely to have reciprocal functional connections to more posterior regions in the word and letter processing network.     

Pure agraphia after infarction in the superior and middle portions of the left precentral gyrus: Dissociation between Kanji and Kana

The present study describes a Japanese patient with pure agraphia displaying differential disturbances in processing Kanji (morphogram) and Kana (syllabogram) letters after an infarction in the middle and superior portions of the left precentral gyrus. Kana errors reflected the patient’s difficulty with retrieving both motor and visual letter images, whereas Kanji errors included partial letter stroke omissions or additions. This present case suggests that differences in writing disturbances between Kana and Kanji letters are caused by a differential dependency on letter motor images.     

Executive functions processed in the frontal and lateral temporal cortices: Intracerebral study

OBJECTIVE: The study was designed to investigate the neurocognitive network in the frontal and lateral temporal cortices that is activated by the complex cognitive visuomotor tasks of letter writing. METHODS: Eight epilepsy surgery candidates with implanted intracerebral depth electrodes performed two tasks involving the writing of single letters. The first task consisted of copying letters. In the second task, the patients were requested to write any other letter. The cognitive load of the second task was increased mainly by larger involvement of the executive functions. The task-related ERD/ERS of the alpha, beta and gamma rhythms was studied. RESULTS: The alpha and beta ERD as the activational correlate of writing of single letters was found in the sensorimotor cortex, anterior cingulate, premotor, parietal cortices, SMA and the temporal pole. The alpha and beta ERD linked to the increased cognitive load was present moreover in the dorsolateral and ventrolateral prefrontal cortex, orbitofrontal cortex and surprisingly also the temporal neocortex. Gamma ERS was detected mostly in the left motor cortex. CONCLUSIONS: Particularly the temporal neocortex was activated by the increased cognitive load. SIGNIFICANCE: The lateral temporal cortex together with frontal areas forms a cognitive network processing executive functions.     

Dual neural circuit model of reading and writing

In the hypothetical neural circuit model of reading and writing that was initially proposed by Dejerine and subsequently confirmed by Geschwind, the left angular gyrus was considered as a unique center for processing letters. Japanese investigators, however, have repeatedly pointed out that this angular gyrus model cannot fully explain the disturbances observed in reading and writing Kanji letters in Japanese patients with various types of alexia with or without agraphia. In 1982, I proposed a dual neural circuit model of reading and writing Japanese on the basis of neuropsychological studies on the various types of alexia with or without agraphia without aphasia. This dual neural circuit model proposes that apart from the left angular gyrus which was thought to be a node for phonological processing of letters, the left posterior inferior temporal area, also acts as a node for semantic processing of letters. Further investigations using O15-PET activation on normal subjects revealed that the left middle occipital gyrus (area 19 of Brodmann) and the posterior portion of the left inferior temporal gyrus (area 37 of Brodmann) are the cortical areas responsible for reading Japanese letters; the former serving for phonological reading and the latter for semantic reading. This duality of the neural circuit in processing letters was later applied to explain disturbances in reading English, and was finally accepted as a valid model for other alphabetic letter systems too.     

What differs in visual recognition of handwritten vs. printed letters? An fMRI study

In models of letter recognition, handwritten letters are considered as a particular font exemplar, not qualitatively different in their processing from printed letters. Yet, some data suggest that recognizing handwritten letters might rely on distinct processes, possibly related to motor knowledge. We applied functional magnetic resonance imaging to compare the neural correlates of perceiving handwritten letters vs. standard printed letters. Statistical analysis circumscribed to frontal brain regions involved in hand‐movement triggering and execution showed that processing of handwritten letters is supported by a stronger activation of the left primary motor cortex and the supplementary motor area. At the whole‐brain level, additional differences between handwritten and printed letters were observed in the right superior frontal, middle occipital, and parahippocampal gyri, and in the left inferior precentral and the fusiform gyri. The results are suggested to indicate embodiment of the visual perception of handwritten letters. Hum Brain Mapp, 2011. © 2010 Wiley‐Liss, Inc.     

Different brain circuits underlie motor and perceptual representations of temporal intervals

In everyday life, temporal information is used for both perception and action, but whether these two functions reflect the operation of similar or different neural circuits is unclear. We used functional magnetic resonance imaging to investigate the neural correlates of processing temporal information when either a motor or a perceptual representation is used. Participants viewed two identical sequences of visual stimuli and used the information differently to perform either a temporal reproduction or a temporal estimation task. By comparing brain activity evoked by these tasks and control conditions, we explored commonalities and differences in brain areas involved in reproduction and estimation of temporal intervals. The basal ganglia and the cerebellum were commonly active in both temporal tasks, consistent with suggestions that perception and production of time are subserved by the same mechanisms. However, only in the reproduction task was activity observed in a wider cortical network including the right pre-SMA, left middle frontal gyrus, left premotor cortex, with a more reliable activity in the right inferior parietal cortex, left fusiform gyrus, and the right extrastriate visual area V5/MT. Our findings point to a role for the parietal cortex as an interface between sensory and motor processes and suggest that it may be a key node in translation of temporal information into action. Furthermore, we discuss the potential importance of the extrastriate cortex in processing visual time in the context of recent findings.     

A case of pure agraphia with a deficit of drawing]

We examined a drawing ability in a case with pure agraphia. The patient, having lesions in the left middle frontal gyrus and inferior parietal lobule, could not write in both dictation and spontaneous writing tasks, with the ability to copy letters being well spared. Correspondingly, he showed a deficit of spontaneous drawing, with the ability to copy complex figures being well spared. These findings suggest that he could have an impairment of retrieving visual image, not restricted to letter image. Although some previous cases with pure agraphia were attributed to an impairment of retrieving visual letter image, drawing ability which seemed to be related to retrieving visual image had not been investigated. In the present case, disability of both writing and drawing could be attributed to a common mechanism, that was an impairment of retrieving visual image. In addition, the patient did not show a significant difference between the performance on Kana writing and on Kanji writing. But it has been said that an impairment of retrieving visual letter image could be related to a deficit of Kanji writing more than of Kana writing. Then, it could be assumed that the mechanism of Kana writing involved not only the pathway depending on graphical motor pattern but one depending on visual image.     

Shared and dissociated cortical regions for object and letter processing

The present study determined the extent to which object and letter recognition recruit similar or dissociated neural resources. Participants passively viewed and silently named line drawings of objects, single letters, and visual noise patterns and centrally fixated an asterisk. We used whole-brain functional MRI and a very conservative approach to hypothesis testing that distinguished among brain regions that were selectively activated by different experimental conditions and those that were conjointly activated. The left fusiform gyrus (BA 19 & 37) and left inferior frontal cortex BA(44/6) showed a greater degree of conjoined activation for objects and letters than selective activation for either category, whereas left inferior parietal cortex (BA 40) and the left insula showed a strong letter-selective response. Equal recruitment of left fusiform and inferior frontal regions by objects and letters reflects similar demands on cognitive processing by these two categories and argues against category-specific modules in these regions. However, cortical systems for object and letter processing are not completely shared given the exclusive activation of left inferior parietal cortex by letters.     

The neural substrates of writing: A functional magnetic resonance imaging study

Background: Hypotheses regarding the neural substrates of writing have been derived from the study of individuals with acquired agraphia. Functional neuroimaging offers another methodology to test these hypotheses in neurologically intact individuals. Aims: This study was designed to identify possible neural substrates for the linguistic and motor components of writing in normal English-speaking individuals. Methods & Procedures: Functional magnetic resonance imaging was used with 12 adults to examine activation associated with generative writing of words from semantic categories contrasted with writing letters of the alphabet and drawing circles. In addition, the generative writing condition was contrasted with a subvocal generative naming condition. Outcomes & Results: Semantically guided retrieval of orthographic word forms for the generative writing condition revealed activation in the left inferior and dorsolateral prefrontal cortex, as well as the left posterior inferior temporal lobe (BA 37). However, no activation was detected in the left angular gyrus (BA 39). The motor components of writing were associated with activation in left fronto-parietal cortex including the region of the intraparietal sulcus, superior parietal lobule, dorsolateral and medial premotor cortex, and sensorimotor areas for the hand. Conclusions: These observations suggest an important role of the left posterior inferior temporal cortex in lexical-orthographic processing and fail to support the long-held notion that the dominant angular gyrus is the storage site for orthographic representations of familiar words. Our findings also demonstrate the involvement of left superior parietal and frontal premotor regions in translating orthographic information into appropriate hand movements.     

Examining the neural and cognitive processes that underlie reading through naming speed tasks

We combined fMRI with eye tracking and speech recording to examine the neural and cognitive mechanisms that underlie reading. To simplify the study of the complex processes involved during reading, we used naming speed (NS) tasks (also known as rapid automatized naming or RAN) as a focus for this study, in which average reading right‐handed adults named sets of stimuli (letters or objects) as quickly and accurately as possible. Due to the possibility of spoken output during fMRI studies creating motion artifacts, we employed both an overt session and a covert session. When comparing the two sessions, there were no significant differences in behavioral performance, sensorimotor activation (except for regions involved in the motor aspects of speech production) or activation in regions within the left‐hemisphere‐dominant neural reading network. This established that differences found between the tasks within the reading network were not attributed to speech production motion artifacts or sensorimotor processes. Both behavioral and neuroimaging measures showed that letter naming was a more automatic and efficient task than object naming. Furthermore, specific manipulations to the NS tasks to make the stimuli more visually and/or phonologically similar differentially activated the reading network in the left hemisphere associated with phonological, orthographic and orthographic‐to‐phonological processing, but not articulatory/motor processing related to speech production. These findings further our understanding of the underlying neural processes that support reading by examining how activation within the reading network differs with both task performance and task characteristics.     

Neural networks for internal reading and visual imagery of reading: a PET study

Regional cerebral blood flow (rCBF) measurements with positron emission tomography (PET) were made on 10 volunteers in rest condition as well as while the subjects, with closed eyes, (i) internally listed the letters of the alphabet and cited the first verse of the Hungarian national anthem, (ii) visualised the capital letters of the alphabet, and (iii) visualised the capital letters of the first verse of the Hungarian national anthem. Significant changes in rCBF indicated various networks of cortical neuronal populations active during the tasks. Internal listing, as compared to the rest condition, activated the left precentral gyrus. Visualising the letters of the alphabet, when compared to the rest condition, activated a cortical network comprising fields along the banks of the left and right intraparietal sulci, the left medial frontal, precentral and occipital sulci, and the right superior frontal gyrus. Visualising the letters of the anthem, when compared to the rest condition, activated a cortical network comprising fields along the banks of the left and right intraparietal sulci, the left medial and inferior frontal gyri, and the right anterior cingulate gyrus. Contrasting the two visualisation tasks revealed task specific activation in the left lateral occipital gyrus (alphabet vs. anthem visualisation) and in the left anterior cingulate gyrus (anthem vs. alphabet visualisation). The data indicate that visual imagery of letters of the alphabet or a text engages a widespread network of cortical fields in the visual association cortices and the frontal cortex, without the engagement of the primary (V1) and secondary (V2) visual cortical areas. This finding supports the hypothesis that neuronal populations engaged by visual imagery and visual perception only partially overlap. The networks, activated in the visualisation tasks, have a core which is identical in the different visualisation tasks. The core network is complemented in a task-specific manner by the recruitment of additional cortical neuronal populations.     

The anterior superior parietal lobule and its interactions with language and motor areas during writing

Past neuroimaging studies of writing demonstrate activation foci in several regions of the posterior parietal cortex (PPC). The present study aimed to dissociate the role of the superior parietal lobule (SPL) from the other PPC regions using functional magnetic resonance imaging (fMRI) and functional connectivity. First, parametric event-related fMRI permitted the categorical comparison of experimental writing conditions with control conditions that were carefully chosen to match the experimental conditions in terms of variables extraneous to the motor act of writing, such as visual stimulus characteristics, and generating and retrieving verbal information. A selective focus of increased activation in the PPC restricted to the rostral part of the SPL (area PE) in the left hemisphere was demonstrated. Subsequently, functional connectivity analysis showed that area PE flexibly interacts with different language areas depending on the linguistic demands of the writing task. Activity in area PE correlates with the left angular gyrus, a region implicated in reading, when the writing is in response to words that are read; in sharp contrast, when the writing is in response to pictured objects, then area PE correlates with the supramarginal gyrus, a region involved in the articulatory and phonological loop, as well as with prefrontal regions that are involved in the retrieval and selection of semantic information. The results suggest that area PE serves as a critical interface between posterior cortical regions in the left hemisphere involved in language processing and the central motor and sensory regions that are directly involved in the control of movement.     

Influence of motor disorders on the visual perception of human movements in a case of peripheral dysgraphia

We report the case of a 71 year-old female patient (DC) with a left parietal lesion resulting in a peripheral dysgraphia essentially characterized by difficulties in letter sequences writing. The aim of our experiments was to analyze the influence of motor difficulties on the visual perception of both writing and reaching movements. Results showed a strong link between motor and perceptual performance. For reaching movements, performances in both production and perception tasks conform to the motor principles identified in healthy subjects (Fitts' law and motor anticipation). By contrast, for handwriting movements, DC's productions do not follow the motor principles usually observed in normal subjects (isochrony principle, motor anticipation) and in perception the same results were observed. The motor references used by DC in the visual perception of writing movement were not the laws of movement but rather her own way of writing. Taken together these data strongly suggest that motor competences is involved in the visual perception of human movements. They are discussed in the general framework of the simulation theory.     

A case of pure agraphia due to left parietal lobe infarction]

We reported a case of an 86-year old woman with pure agraphia due to the left parietal lobe infarction. The characteristics of agraphia were as follows. Most errors in Kana and Kanji writing to dictation and copying were no response. She was able to write only numerals from 1 to 12 precisely. Most errors in numerals were substitution. One unrecognizable numeral was found. She succeeded in pointing to nine among ten single Kana letter named by the examiner with the systematic table of the Japanese syllabary, but missed in pointing to Kana words. It took more time for the patient to point to single Kana letter than the control. Magnetic resonance imaging showed a cerebral infarction in the left parietal lobe which included a part of superior parietal lobule and supramarginal gyrus. We classified pure agraphia with parietal lobe infarction into two types in our previous report. In one type (type 1), letters in writing are poorly formed, but the ability to make words with the methods other than writing are reserved. The only deficit of graphic motor pattern could cause Type 1 agraphia. In another type (type 2), letters in writing were well-formed, but spelling with anagram or typing was disturbed. The deficits of writing process other than graphic motor pattern could cause Type 2 agraphia. This typing seems to be effective not only in Kana but also in Kanji. In this report, we investigated the differences of lesion between two types out of some references. Type1 agraphia seems related to lesion of left superior parietal lobule, while Type 2 agraphia seems related to lesion of left supramarginal gyrus. This case had the features of type 2 agraphia at least, and the compatible lesions.     

Different neural substrates for Kanji and Kana writing: a PET study

To investigate the neural substrate underlying the mechanisms of Kanji and Kana writing, we conducted a PET activation study during mental writing task in eight right-handed normal Japanese subjects. During scans subjects were required to mentally write a Kanji or three Kana letters with their right hand, for each stimulant word presented auditorily. The direct comparisons between Kanji writing and Kana writing revealed that the left posterior inferior temporal gyrus was activated in Kanji writing while the left angular gyrus was activated in Kana writing. In addition, more extensive areas were activated in Kanji writing compared with Kana writing. These results suggest that different respective neural substrates are involved in Kanji and Kana writing respectively.     

Influence of Motor Disorders on the Visual Perception of Human Movements in a Case of Peripheral Dysgraphia

We report the case of a 71 year-old female patient (DC) with a left parietal lesion resulting in a peripheral dysgraphia essentially characterized by difficulties in letter sequences writing. The aim of our experiments was to analyze the influence of motor difficulties on the visual perception of both writing and reaching movements. Results showed a strong link between motor and perceptual performance. For reaching movements, performances in both production and perception tasks conform to the motor principles identified in healthy subjects (Fitts’ law and motor anticipation). By contrast, for handwriting movements, DC’s productions do not follow the motor principles usually observed in normal subjects (isochrony principle, motor anticipation) and in perception the same results were observed. The motor references used by DC in the visual perception of writing movement were not the laws of movement but rather her own way of writing. Taken together these data strongly suggest that motor competences is involved in the visual perception of human movements. They are discussed in the general framework of the simulation theory.     

Selective Kana agraphia: a case report

We present a Japanese man with selective Kana (phonogram) agraphia as a sequela of two cerebral infarctions in a part of the left angular gyrus and its adjoining posterior superior temporal gyrus and the left corona radiata. The agraphia of this patient showed the following features: (1) His writing difficulty was greater for Kana than for Kanji (ideogram) when a word was polysyllabic. (2) Kana errors consisted of perseveration and substitution with another letter, resulting in jargon agraphia. (3) This jargon agraphia often contained a correct number of letters for a target word. Based on these findings, we speculate that the basic defect of this agraphia lies in the process of converting an acoustic word image into a Kana motor grapheme sequence.     

Does visual perception of object afford action? Evidence from a neuroimaging study.

Positron emission tomography (PET) was used to explore the neural correlates of a potential involvement of motor representation during the perception of visually presented objects with different tasks. The main result of this study was that the perception of objects, irrespective of the task (judgement of the vertical orientation, motor imagery, and silent generation of the noun or of the corresponding action verb), versus perception of non-objects, was associated with rCBF increases in a common set of cortical regions. The occipito-temporal junction, the inferior parietal lobule, the SMA-proper, the pars triangularis in the inferior frontal gyrus, the dorsal and ventral precentral gyrus were engaged in the left hemisphere. The ipsilateral cerebellum was also involved. These activations are congruent with the idea of an involvement of motor representation already during the perception of object and thus provide neurophysiological evidence that the perception of objects automatically affords actions that can be made toward them. Besides this common set of cortical areas, each task engaged specific regions.     

Sustained activity within the default mode network during an implicit memory task

Recent neuroimaging studies have shown that several brain regions - namely, the posterior cingulate cortex (PCC), ventral medial prefrontal cortex (vmPFC), and the bilateral angular gyrus - are more active during resting states than during cognitive tasks (i.e., default mode network). Although there is evidence showing that the default mode network is associated with unconscious state, it is unclear whether this network is associated with unconscious processing when normal human subjects perform tasks without awareness. We manipulated the level of conscious processing in normal subjects by asking them to perform an implicit and an explicit memory task, and analyzed signal changes in the default mode network for the stimuli versus baseline in both tasks. The functional magnetic resonance imaging (fMRI) analysis showed that the level of activation in regions within this network during the implicit task was not significantly different from that during the baseline, except in the left angular gyrus and the insula. There was strong deactivation for the explicit task when compared with the implicit task in the default mode regions, except in the left angular gyrus and the left middle temporal gyrus. These data suggest that the activity in the default network is sustained and less disrupted when an implicit memory task is performed, but is suspended when explicit retrieval is required. These results provide evidence that the default mode network is associated with unconscious processing when human subjects perform an implicit memory task.