Päivi Helenius

An MEG Study of Picture Naming

The purpose of this study was to relate a psycholinguistic processing model of picture naming to the dynamics of cortical activation during picture naming. The activation was recorded from eight Dutch subjects with a whole-head neuromagnetometer. The processing model, based on extensive naming latency studies, is a stage model. In preparing a pictures name, the speaker performs a chain of specific operations. They are, in this order, computing the visual percept, activating an appropriate lexical concept, selecting the target word from the mental lexicon, phonological encoding, phonetic encoding, and initiation of articulation. The time windows for each of these operations are reasonably well known and could be related to the peak activity of dipole sources in the individual magnetic response patterns. The analyses showed a clear progression over these time windows from early occipital activation, via parietal and temporal to frontal activation. The major specific findings were that (1) a region in the left posterior temporal lobe, agreeing with the location of Wernickes area, showed prominent activation starting about 200 msec after picture onset and peaking at about 350 msec, (i.e., within the stage of phonological encoding), and (2) a consistent activation was found in the right parietal cortex, peaking at about 230 msec after picture onset, thus preceding and partly overlapping with the left temporal response. An interpretation in terms of the management of visual attention is proposed.

Semantic Cortical Activation in Dyslexic Readers

The combined temporal and spatial resolution of MEG (magnetoencephalography) was used to study whether the same brain areas are similarly engaged in reading comprehension in normal and developmentally dyslexic adults. To extract a semantically sensitive stage of brain activation we manipulated the appropriateness of sentence-ending words to the preceding sentence context. Sentences, presented visually one word at a time, either ended with a word that was (1) expected, (2) semantically appropriate but unexpected, (3) semantically anomalous but sharing the initial letters with the expected word, or (4) both semantically and orthographically inappropriate to the sentence context. In both subject groups all but the highly expected sentence endings evoked strong cortical responses, localized most consistently in the left superior temporal cortex, although additional sources were occasionally found in more posterior parietal and temporal areas and in the right hemisphere. Thus, no significant differences were found in the spatial distribution of brain areas involved in semantic processing between fluent and dyslexic readers. However, both timing and strength of activation clearly differed between the two groups. First, activation sensitivity to word meaning within a sentence context began about 100 msec later in dyslexic than in control subjects. This is likely to result from affected presemantic processing stages in dyslexic readers. Second, the neural responses were significantly weaker in dyslexic than in control subjects, indicating involvement of a smaller or less-synchronous neural population in reading comprehension. Third, in contrast to control subjects, the dyslexic readers showed significantly weaker activation to semantically inappropriate words that began with the same letters as the most expected word than to both orthographically and semantically inappropriate sentence-ending words. Thus, word recognition by the dyslexic group seemed to be qualitatively different: Whereas control subjects perceived words as wholes, dyslexic subjects may have relied on sublexical word recognition and occasionally mistook a correctly beginning word for the one they had expected.

Neural Correlates of Letter-String Length and Lexicality during Reading in a Regular Orthography

Behavioral studies have shown that short letter strings are read faster than long letter-strings and words are read faster than nonwords. Here, we describe the dynamics of letter-string length and lexicality effects at the cortical level, using magnetoencephalography, during a reading task in Finnish with long (eight-letter) and short (four-letter) word/nonword stimuli. Length effects were observed in two spatially and temporally distinct cortical activations: (1) in the occipital cortex at about 100 msec by the strength of activation, regardless of the lexical status of the stimuli, and (2) in the left superior temporal cortex between 200 and 600 msec by the duration of activation, with words showing a smaller effect than nonwords. A significant lexicality effect was also evident in this later activation, with stronger activation and longer duration for nonwords than words. There seem to be no distinct cortical areas for reading words and nonwords. The early length effect is likely to be due to the low-level visual analysis common to all stimulus letter-strings. The later lexicality and length effects apparently reflect converging lexico-semantic and phonological influences, and are discussed in terms of dual-route and single-route connectionist models of reading.

Neurophysiology of fluent and impaired reading: a magnetoencephalographic approach.

This article reviews a series of magnetoencephalographic (MEG) experiments aimed at identifying cortical areas and time windows relevant or even critical for fluent reading. The approach was to compare single-word processing in fluent and dyslexic readers. The activations which differed between the two groups were then studied in more detail to determine their functional roles. In fluent reading, overall visual feature processing occurs about 100 milliseconds (ms) after seeing a word, in the posteromedial extrastriate cortex bilaterally. This activation does not differentiate between letters and symbols. The first reading-specific signal is detected about 150 ms after word onset, when the left inferior occipitotemporal cortex responds preferentially to letter strings. After 200 ms, the left superior temporal cortex, in particular, is engaged in semantic processing of single words and their integration with connected text. While visual feature processing seems to be within normal limits in dyslexic subjects, reading is disrupted during the first 200 ms after seeing a word, at the letter-string specific stage. The subsequent activations are weak and delayed as compared with those in fluent readers. Also presented is a case of deep dyslexia, where the same tools were used to demonstrate that reading comprehension was still subserved by the left hemisphere despite severe damage.

Cortical Effects of Shifting Letter Position in Letter Strings of Varying Length

Neuroimaging and lesion studies suggest that occipito-temporal brain areas play a necessary role in recognizing a wide variety of objects, be they faces, letters, numbers, or household items. However, many questions remain regarding the details of exactly what kinds of information are processed by the occipito-temporal cortex. Here, we address this question with respect to reading. Ten healthy adult subjects performed a single word reading task. We used whole-head magnetoencephalography to measure the spatio-temporal dynamics of brain responses, and investigated their sensitivity to: (1) lexicality (defined here as the difference between words and consonant strings), (2) word length, and (3) variation in letter position. Analysis revealed that midline occipital activity around 100 msec, consistent with low-level visual feature analysis, was insensitive to lexicality and variation in letter position, but was slightly affected by string length. Bilateral occipito-temporal activations around 150 msec were insensitive to lexicality and reacted to word length only in the timing (and not strength) of activation. However, vertical shifts in letter position revealed a hemispheric imbalance: The right hemisphere activation increased with the shifts, whereas the opposite pattern was evident in the left hemisphere. The results are discussed in the light of Caramazza and Hilliss (1990) model of early reading.

Localization of Syntactic and Semantic Brain Responses using Magnetoencephalography

Electrophysiological methods have been used to study the temporal sequence of syntactic and semantic processing during sentence comprehension. Two responses associated with syntactic violations are the left anterior negativity (LAN) and the P600. A response to semantic violation is the N400. Although the sources of the N400 response have been identified in the left (and right) temporal lobe, the neural signatures of the LAN and P600 have not been revealed. The present study used magnetoencephalography to localize sources of syntactic and semantic activation in Finnish sentence reading. Participants were presented with sentences that ended in normally inf lected nouns, nouns in an unacceptable case, verbs instead of nouns, or nouns that were correctly inflected but made no sense in the context. Around 400 msec, semantically anomalous last words evoked strong activation in the left superior temporal lobe with significant activation also for word class errors (N400). Weaker activation was seen for the semantic errors in the right hemisphere. Later, 600-800 msec after word onset, the strongest activation was seen to word class and morphosyntactic errors (P600). Activation was significantly weaker to semantically anomalous and correct words. The P600 syntactic activation was localized to bilateral sources in the temporal lobe, posterior to the N400 sources. The results suggest that the same general region of the superior temporal cortex gives rise to both LAN and N400 with bilateral reactivity to semantic manipulation and a left hemisphere effect to syntactic manipulation. The bilateral P600 response was sensitive to syntactic but not semantic factors.

Abnormal Auditory Cortical Activation in Dyslexia 100 msec after Speech Onset

Reading difficulties are associated with problems in processing and manipulating speech sounds. Dyslexic individuals seem to have, for instance, difficulties in perceiving the length and identity of consonants. Using magnetoencephalography (MEG), we characterized the spatio-temporal pattern of auditory cortical activation in dyslexia evoked by three types of natural bisyllabic pseudowords (/ata/, /atta/, and /a a/), complex nonspeech sound pairs (corresponding to /atta/ and /a a/) and simple 1-kHz tones. The most robust difference between dyslexic and non-reading-impaired adults was seen in the left supratemporal auditory cortex 100 msec after the onset of the vowel /a/. This N100m response was abnormally strong in dyslexic individuals. For the complex nonspeech sounds and tone, the N100m response amplitudes were similar in dyslexic and nonimpaired individuals. The responses evoked by syllable /ta/ of the pseudoword /atta/ also showed modest latency differences between the two subject groups. The responses evoked by the corresponding nonspeech sounds did not differ between the two subject groups. Further, when the initial formant transition, that is, the consonant, was removed from the syllable /ta/, the N100m latency was normal in dyslexic individuals. Thus, it appears that dyslexia is reflected as abnormal activation of the auditory cortex already 100 msec after speech onset, manifested as abnormal response strengths for natural speech and as delays for speech sounds containing rapid frequency transition. These differences between the dyslexic and nonimpaired individuals also imply that the N100m response codes stimulus-specific features likely to be critical for speech perception. Which features of speech (or nonspeech stimuli) are critical in eliciting the abnormally strong N100m response in dyslexic individuals should be resolved in future studies.

Non-impaired auditory phase locking in dyslexic adults

Dyslexic adults have profound difficulties in discriminating rapidly presented sound sequences. To test whether these deficits might be caused by impaired neuronal phase locking to the envelopes of the sound stimuli, 20 normal-reading and 13 dyslexic adults discriminated pitches of pure tones at approximately 1 kHz (producing spectral pitch due to place coding in the cochlea) and of approximately 80 Hz amplitude modulations of white noise (producing periodicity pitch based on temporal information only). We proposed that a specific deficit in phase locking would result in a worse ability to discriminate periodicity than spectral pitch. The dyslexics were significantly less accurate than the control subjects in discriminating both spectral and periodicity pitch stimuli but their performance was not disproportionally worse in the periodicity pitch task. Thus it seems that impaired neuronal phase-locking cannot explain the problems dyslexics face in processing of rapid sound sequences.

Cortical Sequence of Word Perception in Beginning Readers

Efficient analysis of written words in normal reading is likely to reflect use of neural circuits formed by experience during childhood rather than an innate process. We investigated the cortical sequence of word perception in first-graders (7–8 years old), with special emphasis on occipitotemporal cortex in which, in adults, letter-string-sensitive responses are detected at 150 ms after stimulus. To identify neural activation that is sensitive to either the amount of basic visual features or specifically to letter strings, we recorded whole-head magnetoencephalography responses to words embedded in three different levels of noise and to symbol strings. As was shown previously in adults, activation reflecting stimulus nonspecific visual feature analysis was localized to occipital cortex in children. It was followed by letter-string-sensitive activation in the left occipitotemporal cortex and, subsequently, in the temporal cortex. These processing stages were correlated in timing and activation strength. Compared with adults, however, the timing of activation was clearly delayed in children, and the delay was progressively increased from occipital to occipitotemporal and further to temporal areas. This finding is likely to reflect increasing immaturity of the underlying neural generators when advancing from low-level visual analysis to higher-order areas involved in written word perception. When a salient occipitotemporal letter-string-sensitive activation was detected (10 of 18 children), its strength was correlated with phonological skills, in line with the known relevance of phonological awareness in reading acquisition.

Category-specific occipitotemporal activation during face perception in dyslexic individuals: an MEG study

In dyslexia, it is consistently found that letter strings produce an abnormally weak or no response in the left occipitotemporal cortex. Time-sensitive imaging techniques have located this deficit to the category-specific processing stage at about 150 ms after stimulus presentation. The typically reported behavioral impairments in dyslexia suggest that the lack of occipitotemporal activation is specific to reading. It could, however, also reflect a more general dysfunction in the left inferior occipitotemporal cortex or in the time window of category-specific activation (150 to 200 ms). As early cortical processing of faces follows a sequence practically identical to that for letter strings, both in location and in timing, we investigated these possibilities by comparing face-specific occipitotemporal activations in dyslexic and non-reading-impaired subjects. We found that both the stage of general visual feature analysis at about 100 ms and the earliest face-specific activation at about 150 ms were essentially normal in the dyslexic individuals. The present results emphasize the special nature of the occipitotemporal abnormality to letter strings in dyslexia. However, in behavioral tests dyslexic subjects were slower and more error-prone than non-reading-impaired subjects in judging the similarity of faces and geometrical shapes. This effect may be related to reduced activation of the right parietotemporal cortex at about 250 ms after stimulus onset.