During natural speech perception, listeners must track the global speaking rate, that is, the overall rate of incoming linguistic information, as well as transient, local speaking rate variations occurring within the global speaking rate. Here, we address the hypothesis that this tracking mechanism is achieved through coupling of cortical signals to the amplitude envelope of the perceived acoustic speech signals. Cortical signals were recorded with magnetoencephalography (MEG) while participants perceived spontaneously produced speech stimuli at three global speaking rates (slow, normal/habitual, and fast). Inherently to spontaneously produced speech, these stimuli also featured local variations in speaking rate. The coupling between cortical and acoustic speech signals was evaluated using audio–MEG coherence. Modulations in audio–MEG coherence spatially differentiated between tracking of global speaking rate, highlighting the temporal cortex bilaterally and the right parietal cortex, and sensitivity to local speaking rate variations, emphasizing the left parietal cortex. Cortical tuning to the temporal structure of natural connected speech thus seems to require the joint contribution of both auditory and parietal regions. These findings suggest that cortical tuning to speech rhythm operates on two functionally distinct levels: one encoding the global rhythmic structure of speech and the other associated with online, rapidly evolving temporal predictions. Thus, it may be proposed that speech perception is shaped by evolutionary tuning, a preference for certain speaking rates, and predictive tuning, associated with cortical tracking of the constantly changing-rate of linguistic information in a speech stream.

Modern multivariate methods have enabled the application of unsupervised techniques to analyze neurophysiological data without strict adherence to predefined experimental conditions. We demonstrate a multivariate method that leverages priming effects on the evoked potential to perform hierarchical clustering on a set of word stimuli. The current study focuses on the semantic relationships that play a key role in the organization of our mental lexicon of words and concepts. The N400 component of the event-related potential is considered a reliable neurophysiological response that is indicative of whether accessing one concept facilitates subsequent access to another (i.e., one “primes” the other). To further our understanding of the organization of the human mental lexicon, we propose to utilize the N400 component to drive a clustering algorithm that can uncover, given a set of words, which particular subsets of words show mutual priming. Such a scheme requires a reliable measurement of the amplitude of the N400 component without averaging across many trials, which was here achieved using a recently developed multivariate analysis method based on beamforming. We validated our method by demonstrating that it can reliably detect, without any prior information about the nature of the stimuli, a well-known feature of the organization of our semantic memory: the distinction between animate and inanimate concepts. These results motivate further application of our method to data-driven exploration of disputed or unknown relationships between stimuli.

Ten participants learned a miniature language (Anigram), which they later employed to verbally describe a pictured event. Using magnetoencephalography, the cortical dynamics of sentence production in Anigram was compared with that in the native tongue from the preparation phase up to the production of the final word. At the preparation phase, a cartoon image with two animals prompted the participants to plan either the corresponding simple sentence (e.g., “the bear hits the lion”) or a grammar-free list of the two nouns (“the bear, the lion”). For the newly learned language, this stage induced stronger left angular and adjacent inferior parietal activations than for the native language, likely reflecting a higher load on lexical retrieval and STM storage. The preparation phase was followed by a cloze task where the participants were prompted to produce the last word of the sentence or word sequence. Production of the sentence-final word required retrieval of rule-based inflectional morphology and was accompanied by increased activation of the left middle superior temporal cortex that did not differ between the two languages. Activation of the right temporal cortex during the cloze task suggested that this area plays a role in integrating word meanings into the sentence frame. The present results indicate that, after just a few days of exposure, the newly learned language harnesses the neural resources for multiword production much the same way as the native tongue and that the left and right temporal cortices seem to have functionally different roles in this processing.

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.

Neuroimaging and lesion studies suggest that occipitotemporal 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 Hillis's (1990) model of early reading.

We describe a study where a specific treatment method for word-finding difficulty (so-called contextual priming technique, which combines massive repetition priming with semantic priming) was applied with three chronic left hemisphere-damaged aphasics. Both before and after treatment, which focused on naming of a series of pictures, naming-related brain activity was measured by magnetoencephalography (MEG). Due to its excellent temporal resolution and good spatial resolution, we were able to track treatment-induced changes in cortical activity. All three subjects showed improved naming of the trained items. In all subjects, a single source area, located in the left inferior parietal lobe, close to the lesioned area, displayed statistically significant training-induced changes. This effect was of long latency as it started 300–600 msec after picture presentation. The change in activation was specific to training, as it could not be accounted for by variation of cortical dynamics associated with increased proportion of correct answers. Our interpretation is that the training effect reflects more effective phonological encoding and storage of the trained items through the engagement of a left hemispheric word-learning system. This is in line with recent functional imaging studies, which have linked left inferior parietal lobe activity to the phonological storage component of the verbal working memory, as well as with theoretical arguments stating that the primary role of the phonological loop is to acquire new words. Finally, the MEG results showed no evidence of increased right hemisphere participation following training, supporting the view that restoration of language-related networks in the damaged left hemisphere is crucial for anomia recovery.

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.

Magnetoencephalographic (MEG) changes in cortical activity were studied in a chronic Finnish-speaking deep dyslexic patient during single-word and sentence reading. It has been hypothesized that in deep dyslexia, written word recognition and its lexical-semantic analysis are subserved by the intact right hemisphere. However, in our patient, as well as in most nonimpaired readers, lexical-semantic processing as measured by sentence-final semantic-incongruency detection was related to the left superior-temporal cortex activation. Activations around this same cortical area could be identified in single-word reading as well. Another factor relevant to deep dyslexic reading, the morphological complexity of the presented words, was also studied. The effect of morphology was observed only during the preparation for oral output. By performing repeated recordings 1 year apart, we were able to document significant variability in both the spontaneous activity and the evoked responses in the lesioned left hemisphere even though at the behavioural level, the patient's performance was stable. The observed variability emphasizes the importance of estimating consistency of brain activity both within and between measurements in brain-damaged individuals.

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.

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 picture's 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 Wernicke's 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.