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Riitta Salmelin

Publications and source records attributed to Riitta Salmelin.

At least 19 recordsLinked to original sources

Neural representation of language: activation versus long-range connectivity.

Cognitive functions are thought to build on connectivity within large-scale neuronal networks, rather than on strictly localized processes. Yet, present understanding of neural mechanisms of language function, as derived from neuroimaging, is based on mapping brain areas that are more active during specific linguistic tasks than in control conditions. Connectivity can then be evaluated among those areas. However, network nodes should ideally be determined based on their correlated time series of activity. Recent developments in analysis methods now facilitate localization and characterization of functionally connected neural networks directly from real-time magnetoencephalography data. Analysis of long-range connectivity might clarify and expand the view provided by traditional neurophysiological and hemodynamic activation studies. Here, we use silent reading as the example process.

Brain↗

Clinical neurophysiology of language: the MEG approach.

Clinical evaluation of language function and basic neuroscience research into the neurophysiology of language are tied together. Whole-head MEG systems readily facilitate detailed spatiotemporal characterization of language processes. A fair amount of information is available about the cortical sequence of word perception and comprehension in the auditory and visual domain, which can be applied for clinical use. Language production remains, at present, somewhat less well charted. In clinical practice, the most obvious needs are noninvasive evaluation of the language-dominant hemisphere and mapping of areas involved in language performance to assist surgery. Multiple experimental designs and analysis approaches have been proposed for estimation of language lateralization. Some of them have been compared with the invasive Wada test and need to be tested further. Development of approaches for more comprehensive pre-surgical characterization of language cortex should build on basic neuroscience research, making use of parametric designs that allow functional mapping. Studies of the neural basis of developmental and acquired language disorders, such as dyslexia, stuttering, and aphasia can currently be regarded more as clinical or basic neuroscience research rather than as clinical routine. Such investigations may eventually provide tools for development of individually targeted training procedures and their objective evaluation.

Brain↗

Cortical dynamics of visual/semantic vs. phonological analysis in picture confrontation.

Picture naming covers the main stages of word production from concept retrieval to articulation. Cortical correlates of picture naming have been characterized with both haemodynamic and neurophysiological methods but the association of specific activation patterns with the hypothesized processing stages remains elusive. Here we used categorization tasks to selectively highlight different components of picture confrontation, from visual analysis (VIS) to semantic (SEM) and phonological access (PHON), and compared these time courses of activation with that obtained during picture naming (NAM). Brain activity was recorded with whole-head magnetoencephalography (MEG). Following the initially similar activation patterns in occipital and parietal areas, task effects (stronger activation in NAM/PHON than in SEM/VIS) emerged after 300 ms, in the sustained activation of the left posterior temporal and bilateral inferior frontal cortex, apparently reflecting enhancement of phonological and phonetic/articulatory processing.

Adult↗

Phase coupling in a cerebro-cerebellar network at 8-13 Hz during reading.

Words forming a continuous story were presented to 9 subjects at frequencies ranging from 5 to 30 Hz, determined individually to render comprehension easy, effortful, or practically impossible. We identified a left-hemisphere neural network sensitive to reading performance directly from the time courses of activation in the brain, derived from magnetoencephalography data. Regardless of the stimulus rate, communication within the long-range neural network occurred at a frequency of 8-13 Hz. Our coherence-based detection of interconnected nodes reproduced several brain regions that have been previously reported as active in reading tasks, based on traditional contrast estimates. Intriguingly, the face motor cortex and the cerebellum, typically associated with speech production, and the orbitofrontal cortex, linked to visual recognition and working memory, additionally emerged as densely connected components of the network. The left inferior occipitotemporal cortex, involved in early letter-string or word-specific processing, and the cerebellum turned out to be the main forward driving nodes of the network. Synchronization within a subset of nodes formed by the left occipitotemporal, the left superior temporal, and orbitofrontal cortex was increased with the subjects' effort to comprehend the text. Our results link long-range neural synchronization and directionality with cognitive performance.

Adult↗

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.

Adolescent↗

Motor cortex dynamics in visuomotor production of speech and non-speech mouth movements.

We investigated timing and hemispheric balance of motor cortex activation when kinetically similar speech and non-speech mouth movements and sequences of such movements were triggered by visually presented letter- and symbol-strings. As an index of motor cortex activation, we used magnetoencephalographic recording of task-related change of precentral 20 Hz (16-24 Hz) activity. Suppression of the 20 Hz rhythm revealed pre-movement activation in the face representation areas that was tied to visual instruction, not movement onset. The 20 Hz rhythm remained suppressed throughout the preparation and execution of mouth movements and was followed by post-movement rebound. Left hemisphere preceded the right at the onset and offset of the suppression, similarly for isolated and sequential speech and non-speech movements. Pattern of task-related change in 20 Hz activity was otherwise symmetrical. In the face areas, the overall modulation of 20 Hz activity increased with sequence length and motor demands. Hand representation areas showed also weak reactivity, with systematically larger modulation of 20 Hz activity for non-speech than speech movements. Our results suggest an active role for the motor cortex in cognitive control of visually triggered mouth movements, not limited to movement execution.

Adult↗

Right rolandic activation during speech perception in stutterers: a MEG study.

The focus of our magnetoencephalographic (MEG) study was to obtain further insight into the neuronal organization of language processing in stutterers. We recorded neuronal activity of 10 male developmental stutterers and 10 male controls, while they listened to pure tones, to words in order to repeat them, and to sentences in order to either repeat or transform them into passive form. Stimulation with pure tones resulted in similar activation patterns in the two groups, but differences emerged in the more complex auditory language tasks. In the stutterers, the left inferior frontal cortex was activated for a short while from 95 to 145 ms after sentence onset, which was not evident in the controls nor in either group during the word task. In both subject groups, the left rolandic area was activated when listening to the speech stimuli, but in the stutterers, there was an additional activation of the right rolandic area from 315 ms onwards, which was more pronounced in the sentence than word task. Activation of areas typically associated with language production was thus observed also during speech perception both in controls and in stutterers. Previous research on speech production in stutterers has found abnormalities in both the amount and timing of activation in these areas. The present data suggest that activation in the left inferior frontal and right rolandic areas in stutterers differs from that in controls also during speech perception.

Adolescent↗

Time course of top-down and bottom-up influences on syllable processing in the auditory cortex.

In speech perception, extraction of meaning from complex streams of sounds is surprisingly fast and efficient. By tracking the neural time course of syllable processing with magnetoencephalography we show that this continuous construction of meaning-based representations is aided by both top-down (context-based) expectations and bottom-up (acoustic-phonetic) cues in the speech signal. Syllables elicited a sustained response at 200-600 ms (N400m) which became most similar to that evoked by words when the expectation for meaningful speech was increased by presenting the syllables among words and sentences or using sentence-initial syllables. This word-like cortical processing of meaningless syllables emerged at the build-up of the N400m response, 200-300 ms after speech onset, during the transition from perceptual to lexical-semantic analysis. These findings show that the efficiency of meaning-based analysis of speech is subserved by a cortical system finely tuned to lexically relevant acoustic-phonetic and contextual cues.

Adult↗

Cortical differentiation of speech and nonspeech sounds at 100 ms: implications for dyslexia.

Neurophysiological measures indicate cortical sensitivity to speech sounds by 150 ms after stimulus onset. In this time window dyslexic subjects start to show abnormal cortical processing. We investigated whether phonetic analysis is reflected in the robust auditory cortical activation at approximately 100 ms (N100m), and whether dyslexic subjects show abnormal N100m responses to speech or nonspeech sounds. We used magnetoencephalography to record auditory responses of 10 normally reading and 10 dyslexic adults. The speech stimuli were synthetic Finnish speech sounds (/a/, /u/, /pa/, /ka/). The nonspeech stimuli were complex nonspeech sounds and simple sine wave tones, composed of the F1+F2+F3 and F2 formant frequencies of the speech sounds, respectively. All sounds evoked a prominent N100m response in the bilateral auditory cortices. The N100m activation was stronger to speech than nonspeech sounds in the left but not in the right auditory cortex, in both subject groups. The leftward shift of hemispheric balance for speech sounds is likely to reflect analysis at the phonetic level. In dyslexic subjects the overall interhemispheric amplitude balance and timing were altered for all sound types alike. Dyslexic individuals thus seem to have an unusual cortical organization of general auditory processing in the time window of speech-sensitive analysis.

Acoustic Stimulation↗

Activation of the human sensorimotor cortex during error-related processing: a magnetoencephalography study.

We studied error-related processing using magnetoencephalography (MEG). Previous event-related potential studies have documented error negativity or error-related negativity after incorrect responses, with a suggested source in the anterior cingulate cortex or supplementary motor area. We compared activation elicited by correct and incorrect trials using auditory and visual choice-reaction time tasks. Source areas showing different activation patterns in correct and error conditions were mainly located in sensorimotor areas, both ipsi- and contralateral to the response, suggesting that activation of sensorimotor circuits accompanies error processing. Additional activation at various other locations suggests a distributed network of brain regions active during error-related processing. Activation specific to incorrect trials tended to occur later in MEG than EEG data, possibly indicating that EEG and MEG detect different neural networks involved in error-related processes.

Acoustic Stimulation↗

Learning new names for new objects: cortical effects as measured by magnetoencephalography.

We tracked the evolvement of naming-related cortical dynamics with magnetoencephalography when five normal adults successfully learned names and/or meanings of unfamiliar objects. In all subjects, the learning of new names was associated with pronounced cortical effects. The learning effect was of long latency and emerged as a change of activation in the same cortical network that was active during naming of familiar items. In four out of five subjects, the cortical learning effect occurred in the inferior parietal lobe. In three of these subjects, the cortical effect was left-sided. These results suggest that the inferior parietal lobe plays an important role in the acquisition of novel words, presumably as a part of working memory systems.

Adult↗

Cortical activation during a spatiotemporal tactile comparison task.

Tactile sensory memory is needed to infer shape or motion from the spatiotemporal pattern of sensory input during manual exploration. Here we applied triplets of pressure pulses to the fingertips of subjects who were asked to respond when successive triplets were the same (COMPARE task) or when a particular stimulus was included in a triplet (CONTROL task). Stimulus sequences (30 s) alternated with rest blocks (30 s) and functional magnetic resonance images (fMRIs) were acquired in a 1.5-T scanner. During the COMPARE task, we found enhanced activation in inferior parietal cortex, supplementary motor area (SMA), and right dorsolateral prefrontal cortex (DLPFC). Activation of DLPFC is likely to be related to the attempt to memorize the stimulus sequences and activations of SMA and inferior parietal cortex to the analysis of temporospatial tactile patterns and, more generally, to guidance of haptic exploration. In addition, task-specific activation was seen in anterior cingulate gyrus, possibly related to the high mental effort required by the comparison task. Our rhythmic tactile stimulus as such, without any task-specific enhancement, activated also left cerebellum and (mainly left) putamen, supporting the idea that these structures are related to perception of temporal order of tactile stimuli.

Adult↗

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 Hillis's (1990) model of early reading.

Adult↗

Adult brain plasticity elicited by anomia treatment.

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.

Aphasia↗

Hemispheric balance in processing attended and non-attended vowels and complex tones.

We compared cortical processing of attended and non-attended vowels and complex tones, using a whole-head neuromagnetometer, to test for possible hemispheric differences. Stimuli included vowels [a] and [i], spoken by two female Finnish speakers, and two complex tones, each with two pure tone components corresponding to the first and second formant frequencies (F1-F2) of the vowels spoken by speaker 1. Sequences including both vowels and complex tones were presented to eight Finnish males during passive and active (phoneme/speaker/complex tone identification) listening. Sequences including only vowels were presented to five of the subjects during passive listening and during a phoneme identification task. The vowel [i] spoken by speaker 1 and the corresponding complex tone were frequent, non-target stimuli. Responses evoked by these frequent stimuli were analyzed. Cortical activation at approximately 100 ms was stronger for the complex tone than the vowel in the right hemisphere (RH). Responses were similar during active and passive listening. Hemispheric balance remained the same when the vowel was presented in sequences including only vowels. The reduction of RH activation for vowels as compared with complex tones indicates a relative increase of left hemisphere involvement, possibly reflecting a shift towards more language-specific processing.

Acoustic Stimulation↗

Naming actions and objects: cortical dynamics in healthy adults and in an anomic patient with a dissociation in action/object naming.

Neuropsychological studies have demonstrated that the production of nouns and verbs can be dissociated in aphasia. These reports have been taken as evidence for separate representations of nouns and verbs in the human brain. We used whole-head magnetoencephalography to record cortical dynamics of action and object naming in 10 healthy adults and in 1 anomic patient with superior naming of verbs compared with nouns due to a left posterior parietal lesion. A single set of 100 line drawings was used for both action and object naming. In normal subjects, the activation sequences in action and object naming were essentially identical, advancing from the occipital to posterior temporoparietal and further to the left frontal cortex, without consistent involvement of the classical left inferior frontal (Broca) and temporal (Wernicke) language areas. In the anomic patient, pronounced differences between action and object naming emerged in the left hemisphere. The activation sequence was disrupted at the level of the damaged parietal cortex and did not reach the left frontal cortex even in the relatively easier action naming. The more severely impaired object naming was associated with exceptionally strong and early activation of the left inferior frontal cortex (Broca) and subsequent pronounced activation of the left middle temporal cortex, silent in action naming. Verb and noun retrieval thus utilized a spatiotemporally similar neuronal network in healthy individuals. A clear dissociation in cortical correlates of verb and noun retrieval only became evident in our anomic patient, in whom damage to the language network has resulted in disproportionately worse performance in object than action naming.

Adult↗

Comparison of BOLD fMRI and MEG characteristics to vibrotactile stimulation.

The characteristics of blood oxygenation level-dependent (BOLD) fMRI and magnetoencephalographic (MEG) responses to vibrotactile stimuli in humans were studied and compared. The stimuli, presented with interstimulus intervals (ISIs) ranging from 1 to 5 s, yielded highly reproducible MEG responses, with current dipoles in the primary somatosensory (SI) cortex in all subjects. BOLD fMRI responses to similar stimuli showed substantial intrasubject variation in the activation sites around the SI cortex. BOLD responses were detected in all subjects in the secondary somatosensory (SII) cortices as well, with comparable BOLD response amplitudes to those in the SI cortex. Current dipoles, used to model the MEG signals, were stronger at longer ISIs than shorter ISIs. The BOLD response amplitudes did not show a similar dependence on ISI, but the activated brain area was larger when longer ISIs or longer stimuli were applied. Our results support the view that combined use of brain mapping methods provides complementary information and should be considered in functional brain examinations.

Adult↗

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.

Acoustic Stimulation↗