The present fMRI study investigated the effects of word-likeness of visual and auditory stimuli on activity along the ventral visual stream. In the context of a one-back task, we presented visual and auditory words, pseudowords, and artificial stimuli (i.e., false-fonts and reversed-speech, respectively). Main findings were regionally specific effects of word-likeness on activation in a left ventral occipitotemporal region corresponding to the classic localization of the Visual Word Form Area (VWFA). Specifically, we found an inverse word-likeness effect for the visual stimuli in the form of decreased activation for words compared to pseudowords which, in turn, elicited decreased activation compared to the artificial stimuli. For the auditory stimuli, we found positive word-likeness effects as both words and pseudowords elicited more activation than the artificial stimuli. This resulted from a marked deactivation in response to the artificial stimuli and no such deactivation for words and pseudowords. We suggest that the opposite effects of visual and auditory word-likeness on VWFA activation can be explained by assuming the involvement of visual orthographic memory representations. For the visual stimuli, these representations reduce the coding effort as a function of word-likeness. This results in highest activation to the artificial stimuli and least activation to words for which corresponding representations exist. The positive auditory word-likeness effects may result from activation of orthographic information associated with the auditory words and pseudowords. The view that the VWFA has a primarily visual function is supported by our findings of high activation to the visual artificial stimuli (which have no phonological or semantic associations) and deactivation to the auditory artificial stimuli. According to the phenomenon of cross-modal sensory suppression such deactivations during demanding auditory processing are expected in visual regions.
fMRI; neuroimaging; one-back task; word-likeness; word processing; VWFA; orthographic representations
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 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).
The importance of the left occipitotemporal cortex for visual word processing is highlighted by numerous functional neuroimaging studies, but the precise function of the Visual Word Form Area (VWFA) in this brain region is still under debate. The present fMRI study varied orthographic familiarity independent from phonological-semantic familiarity by presenting orthographically familiar and orthographically unfamiliar forms (pseudohomophones) of the same words in a phonological lexical decision task. Consistent with orthographic word recognition in the VWFA, we found lower activation for familiar compared to unfamiliar forms, but no difference between pseudohomophones and pseudowords. This orthographic familiarity effect in the VWFA differed from the phonological familiarity effect in left frontal regions, where phonologically unfamiliar pseudowords led to higher activation than phonologically familiar pseudohomophones. We suggest that the VWFA not only computes letter string representations but also hosts word specific orthographic representations. These representations function as recognition units with the effect that letter strings, which readily match with stored representations lead to less activation than letter strings which do not.
Functional MRI; orthographic word recognition; visual word processing; occipitotemporal cortex; reading
Based on our previous work, we expected the Visual Word Form Area (VWFA) in the left ventral visual pathway to be engaged by both whole-word recognition and by serial sublexical coding of letter strings. To examine this double function, a phonological lexical decision task (i.e., “Does xxx sound like an existing word?”) presented short and long letter strings of words, pseudohomophones, and pseudowords (e.g., Taxi, Taksi and Tazi). Main findings were that the length effect for words was limited to occipital regions and absent in the VWFA. In contrast, a marked length effect for pseudowords was found throughout the ventral visual pathway including the VWFA, as well as in regions presumably engaged by visual attention and silent-articulatory processes. The length by lexicality interaction on brain activation corresponds to well-established behavioral findings of a length by lexicality interaction on naming latencies and speaks for the engagement of the VWFA by both lexical and sublexical processes.
This study used foci from 17 original studies on functional abnormalities in the dyslexic brain to identify brain regions with consistent under- or overactivation. Studies were included when reading or reading-related tasks were performed on visually presented stimuli and when results reported coordinates for group differences. Activation Likelihood Estimation (ALE) was used for quantification. Maxima of underactivation were found in inferior parietal, superior temporal, middle and inferior temporal and fusiform regions of the left hemisphere. With respect to left frontal abnormalities we found underactivation in the inferior frontal gyrus to be accompanied by overactivation in the primary motor cortex and the anterior insula. Tentative functional interpretations of the activation abnormalities are provided.
Dyslexia; Reading; Magnetic Resonance Imaging; Positron-Emission Tomography; Cerebral Cortex
Impairments of the lexical and the nonlexical reading route were examined for German-speaking dyslexic readers by measuring accuracy and speed of phonological and orthographic lexical decisions. Different from English-based findings, we found little difficulty with the phonological distinction between pseudohomophones and nonwords, but a major difficulty with the orthographic distinction between words and pseudohomophones. Subtyping identified pure surface dyslexia cases but no case of pure phonological dyslexia. Dyslexic speed impairments were traced to three loci in the dual-route model: an impoverished orthographic lexicon, slow access from orthographic to phonological lexicon entries (lexical route) and from graphemes to phonemes (nonlexical route). A review of distal cognitive deficits suggested that the orthographic lexicon is affected by phonological deficits and that the slow functioning of the lexical and the nonlexical route reflects a general visual-verbal speed impairment and not a purely visual-attentional deficit.
dyslexia; phonological lexical decision; orthographic lexical decision; reading speed impairment; dual-route model
This fMRI study contrasted case-deviant and letter-deviant forms with familiar forms of the same phonological words (e.g., TaXi and Taksi vs. Taxi) and found, that both types of deviance led to increased activation in a left occipitotemporal region corresponding to the Visual Word Form Area. Case-deviant items, in addition, led to increased activation in a right occipitotemporal region and in a left occipital and a left posterior occipitotemporal region, possibly reflecting the increased demands on letter form coding. For letter-deviant items, in addition to the increased left occipitotemporal activation, a main finding was increased activation primarily in extended left frontal regions, possibly reflecting sublexically mediated access to word phonology. These findings are consistent with general features of cognitive dual-route models of visual word processing. Furthermore, they add support to the main feature of Dehaene et al.’s (2005) neural model of early stages of visual word processing . However, the increased activation found for case-deviant items in the VWFA cannot be immediately reconciled with the assumption of completely abstract case-independent orthographic word codes in the VWFA.
Functional MRI; visual word recognition; occipitotemporal cortex; visual word form area; orthographic processing
This study assessed eye movement abnormalities of adolescent dyslexic readers and interpreted the findings by linking the dual-route model of single word reading with the E-Z Reader model of eye movement control during silent sentence reading. A dysfunction of the lexical route was assumed to account for a reduced number of words which received only a single fixation or which were skipped and for the increased number of words with multiple fixations and a marked effect of word length on gaze duration. This pattern was interpreted as a frequent failure of orthographic whole-word recognition (based on orthographic lexicon entries) and on reliance on serial sublexical processing instead. Inefficiency of the lexical route was inferred from prolonged gaze durations for singly fixated words. These findings were related to the E-Z Reader model of eye movement control. Slow activation of word phonology accounted for the low skipping rate of dyslexic readers. Frequent reliance on sublexical decoding was inferred from a tendency to fixate word beginnings and from short forward saccades. Overall, the linkage of the dual-route model of single word reading and a model of eye movement control led to a useful framework for understanding eye movement abnormalities of dyslexic readers.
Developmental dyslexia; Dual-route model of reading; Eye movement control; E-Z Reader; Lexical route; Sublexical route
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).
Developmental dyslexia; fMRI; Reading; Phonological lexical decision; Dual-route
In two previous studies we assessed a diffculty of dyslexic readers with letter string processing by using variants of the partial report paradigm, e.g., Averbach and Coriell [Averbach, E., & Coriell, A. S. (1961). Short-term memory in vision. Bell Systems Technical Journal, 40, 309-328] which requires report of a letter name in response to a position cue. The poor dyslexic performance was interpreted as evidence for a visual-attentional deficit of dyslexic readers. In the present study, we avoided verbal report by using a task which only required the detection of predefined targets (letters or pseudoletters) in strings. On this purely visual task, the dyslexic readers did not differ from non-impaired readers. This finding speaks against a basic visual-attentional deficit; rather it suggests that the dyslexic deficit on partial report paradigms stems from a problem in establishing a string representation which includes position and name codes.
Developmental dyslexia; Target detection; Visual attention; String processing
We used fMRI to examine functional brain abnormalities of German-speaking dyslexics who suffer from slow effortful reading but not from a reading accuracy problem. Similar to acquired cases of letter-by-letter reading, the developmental cases exhibited an abnormal strong effect of length (i.e., number of letters) on response time for words and pseudowords.
Corresponding to lesions of left occipito-temporal (OT) regions in acquired cases, we found a dysfunction of this region in our developmental cases who failed to exhibit responsiveness of left OT regions to the length of words and pseudowords. This abnormality in the left OT cortex was accompanied by absent responsiveness to increased sublexical reading demands in phonological inferior frontal gyrus (IFG) regions. Interestingly, there was no abnormality in the left superior temporal cortex which—corresponding to the onological deficit explanation—is considered to be the prime locus of the reading difficulties of developmental dyslexia cases.
The present functional imaging results suggest that developmental dyslexia similar to acquired letter-by-letter reading is due to a primary dysfunction of left OT regions.