Kuniyoshi L. Sakai

Preferential activation of different I waves by transcranial magnetic stimulation with a figure-of-eight-shaped coil.

Transcranial magnetic stimulation (TMS) over the human primary motor cortex (M1) evokes motor responses in the contralateral limb muscles. The latencies and amplitudes of those responses depend on the direction of induced current in the brain by the stimuli (Mills et al. 1992, Werhahn et al. 1994). This observation suggests that different neural elements might be activated by the differently directed induced currents. Using a figure-of-eight-shaped coil, which induces current with a certain direction, we analyzed the effect of direction of stimulating current on the latencies of responses to TMS in normal subjects. The latencies were measured from surface electromyographic responses of the first dorsal interosseous muscles and the peaks in the peristimulus time histograms (PSTHs) of single motor units from the same muscles. The coil was placed over the M1, with eight different directions each separated by 45°. Stimulus intensity was adjusted just above the motor threshold while subjects made a weak tonic voluntary contraction, so that we can analyse the most readily elicited descending volley in the pyramidal tracts. In most subjects, TMS with medially and anteriorly directed current in the brain produced responses or a peak that occurred some 1.5 ms later than those to anodal electrical stimulation. In contrast, TMS with laterally and posteriorly directed current produced responses or a peak that occurred about 4.5 ms later. There was a single peak in most of PSTHs under the above stimulation condition, whereas there were occasionally two peaks under the transitional current directions between the above two groups. These results suggest that TMS with medially and anteriorly directed current in the brain readily elicits I1 waves, whereas that with laterally and posteriorly directed current preferentially elicits I3 waves. Functional magnetic resonance imaging studies indicated that this direction was related to the course of the central sulcus. TMS with induced current flowing forward relative to the central sulcus preferentially elicited I1 waves and that flowing backward elicited I3 waves. Our finding of the dependence of preferentially activated I waves on the current direction in the brain suggests that different sets of cortical neurons are responsible for different I waves, and are contrarily oriented. The present method using a figure-of-eight-shaped coil must enable us to study physiological characteristics of each I wave separately and, possibly, analyse different neural elements in M1, since it activates a certain I wave selectively without D waves or other I waves.

Temporal cortex activation during speech recognition: an optical topography study.

Cortical activity during speech recognition was examined using optical topography (OT), a recently developed non-invasive technique. To assess relative changes in hemoglobin oxygenation, local changes in near-infrared light absorption were measured simultaneously from 44 points in both hemispheres. A dichotic listening paradigm was used in this experiment, in which target stimuli and non-target stimuli were presented to different ears. Subjects were asked to track targets and to press a button when targets shifted from one ear to the other. We compared three tasks: (i) a control task, in which a tone was used as the target; (ii) a repeat task, in which the target was one repeated sentence; (iii) a story task, in which the targets were continuous sentences of a story. The activity for the story task, compared with the repeat task, was localized in the left superior temporal cortex. Relative to the control task, we observed in this region a larger increase in oxyhemoglobin concentration and a decrease in deoxyhemoglobin concentration in the story task than those in the repeat task. These results suggest that the activity in the left temporal association area reflects the load of auditory, memory, and language information processing.

Brain Activations During Conscious Self- Monitoring of Speech Production With Delayed Auditory Feedback: An fMRI Study

When a speakers voice returns to ones own ears with a 200‐ms delay, the delay causes the speaker to speak less fluently. This phenomenon is called a delayed auditory feedback (DAF) effect. To investigate neural mechanisms of speech processing through the DAF effect, we conducted a functional magnetic resonance imaging (fMRI) experiment, in which we designed a paradigm to explore the conscious overt‐speech processing and the automatic overt‐speech processing separately, while reducing articulatory motion artifacts. The subjects were instructed to (1) read aloud visually presented sentences under real‐time auditory feedback (NORMAL), (2) read aloud rapidly under real‐time auditory feedback (FAST), (3) read aloud slowly under real‐time auditory feedback (SLOW), and (4) read aloud under DAF (DELAY). In the contrasts of DELAY‐NORMAL, DELAY‐FAST, and DELAY‐SLOW, the bilateral superior temporal gyrus (STG), the supramarginal gyrus (SMG), and the middle temporal gyrus (MTG) showed significant activation. Moreover, we found that the STG activation was correlated with the degree of DAF effect for all subjects. Because the temporo‐parietal regions did not show significant activation in the comparisons among NORMAL, FAST, and SLOW conditions, we can exclude the possibility that its activation is due to speech rates or enhanced attention to altered speech sounds. These results suggest that the temporo‐parietal regions function as a conscious self‐monitoring system to support an automatic speech production system. Hum. Brain Mapping 20:22–28, 2003.

Specialization in the Left Prefrontal Cortex for Sentence Comprehension

Using functional magnetic resonance imaging (fMRI), we examined cortical activation under syntactic decision tasks and a short-term memory task for sentences, focusing on essential properties of syntactic processing. By comparing activation in these tasks with a short-term memory task for word lists, we found that two regions in the left prefrontal cortex showed selective activation for syntactic processing: the dorsal prefrontal cortex (DPFC) and the inferior frontal gyrus (IFG). Moreover, the left DPFC showed more prominent activation under the short-term memory task for sentences than that for word lists, which cannot be explained by general cognitive factors such as task difficulty and verbal short-term memory. These results support the proposal of specialized systems for sentence comprehension in the left prefrontal cortex.

Functional mapping of the human colour centre with echo-planar magnetic resonance imaging

Clinical studies of cerebral achromatopsia have suggested a colour centre in the human fusiform gyrus. By using functional magnetic resonance imaging, we examined whether the fusiform gyrus shows activity correlated with the perception of colour. We tested three stimulus conditions in which the subject maintained fixation: (i) a circular array of six coloured circles; (ii) the same as (i) except that each circle is equiluminant grey with its colour counterpart; and (iii) the same as (i) plus a clockwise shift of circles to neighbouring positions every 1 s. After termination of the stimulus, the subject perceived an after-image of circles with complementary colours in (i), but not in (iii). In condition (i), we found a focal signal increase in the posterior part of the fusiform gyrus. In condition (ii), the activation in the same locus during the stimulation period was weaker than that in (i). In condition (iii), the signal intensity after termination of the stimulus was weaker than that in (i). The colour effect and after-effect on activation of the fusiform gyrus observed here suggest its critical role in human colour perception.

Learning Letters in Adulthood: Direct Visualization of Cortical Plasticity for Forming a New Link between Orthography and Phonology

To identify which brain regions in adults show plasticity for learning letters, Hangul letters were experimentally associated with either speech sounds (HS condition) or nonspeech sounds (HN condition) in fMRI sessions over two consecutive days. Selective activations under the HS condition were found in several regions including the left posterior inferior temporal gyrus (PITG) and the parieto-occipital cortex (PO), as compared with activations under a condition for familiar letters and speech sounds, and with those under the HN condition. The left PITG showed a selective activation increase under the HS condition over two days, the degree of which predicted individual performance improvement. Further, functional connectivity between the left PITG and the left PO was selectively enhanced under the HS condition. These results demonstrate that a new link between orthography and phonology is formed by the plasticity of a functional system involving the left PITG in association with the left PO.

From Perception to Sentence Comprehension: The Convergence of Auditory and Visual Information of Language in the Left Inferior Frontal Cortex

We used functional magnetic resonance imaging (fMRI) to characterize cortical activation associated with sentence processing, thereby elucidating where in the brain auditory and visual inputs of words converge during sentence comprehension. Within one scanning session, subjects performed three types of tasks with different linguistic components from perception to sentence comprehension: nonword (N(AV); auditory and visual), phrase (P; either auditory or visual), and sentence (S; either auditory or visual) tasks. In a comparison of the P and N(AV) tasks, the angular and supramarginal gyri showed bilateral activation, whereas the inferior and middle frontal gyri showed left-lateralized activation. A comparison of the S and P tasks, together with a conjunction analysis, revealed a ventral region of the left inferior frontal gyrus (F3t/F3O), which was sentence-processing selective and modality-independent. These results unequivocally demonstrated that the left F3t/F3O is involved in the selection and integration of semantic information that are separable from lexico-semantic processing.

Functional differentiation in the human auditory and language areas revealed by a dichotic listening task.

The human auditory cortex plays a special role in speech recognition. It is therefore necessary to clarify the functional roles of individual auditory areas. We applied functional magnetic resonance imaging (fMRI) to examine cortical responses to speech sounds, which were presented under the dichotic and diotic (binaural) listening conditions. We found two different response patterns in multiple auditory areas and language-related areas. In the auditory cortex, the medial portion of the secondary auditory area (A2), as well as a part of the planum temporale (PT) and the superior temporal gyrus and sulcus (ST), showed greater responses under the dichotic condition than under the diotic condition. This dichotic selectivity may reflect acoustic differences and attention-related factors such as spatial attention and selective attention to targets. In contrast, other parts of the auditory cortex showed comparable responses to the dichotic and diotic conditions. We found similar functional differentiation in the inferior frontal (IF) cortex. These results suggest that multiple auditory and language areas may play a pivotal role in integrating the functional differentiation for speech recognition.

Language-Related Activations in the Left Prefrontal Regions Are Differentially Modulated by Age, Proficiency, and Task Demands

It remains to be elucidated how cortical activations are modulated by factors of age, proficiency, and language task demands when mastering first language (L1) and a second language (L2). Using functional magnetic resonance imaging, we tested subjects aged 13 (the age 13 group) and 19 (the age 19 group), thereby comparing the cortical activations involved in past-tense verb identification with those involved in verb matching. We found that the activation in the dorsal triangular part of the left inferior frontal gyrus (IFG) was lower, corresponding to a higher proficiency in English (L2) in the older subjects, suggesting that the proficiency level plays a major role in the activation of this region during L2 acquisition. Moreover, the lower activation in the triangular and orbital parts of the left IFG (F3t/F3O) for the irregular past tense corresponding to a higher proficiency in L2, together with the nonsignificant activation for the regular past tense when its performance almost reached perfection for age 19, suggests that the modulation of the left F3t/F3O activation reflects language task demands for identifying correct past-tense forms. On the other hand, the left F3t/F3O activation in Japanese (L1) for age 13 was significantly greater than that for age 19, despite the matched performances in L1. These results suggest that the left IFG subserves language-specific functions that are critically required when mastering any language.

Lateralized activation in the inferior frontal cortex during syntactic processing: Event-related optical topography study

Functional imaging with near‐infrared light has the potential to provide novel information that cannot be obtained with other imaging techniques. An event‐related paradigm has not been fully established for studying human cognitive functions with near‐infrared optical imaging. We conducted language experiments to develop an event‐related paradigm with optical topography (OT). We directly compared cortical activation during syntactic and semantic decision tasks, both of which involved error detection in a sentence stimulus that consisted of a noun phrase and a verb. In the syntactic decision task, subjects judged whether the presented sentence is syntactically correct, where the syntactic knowledge about the distinction between transitive and intransitive verbs was required. In the semantic decision task, subjects judged whether the presented sentence is semantically correct, where the lexico‐semantic knowledge about selectional restrictions was indispensable. We found local increases in oxyhemoglobin concentration, which were selectively associated with the syntactic decision task. Activation in the left inferior frontal gyrus was detected when syntactically anomalous sentences were presented, whereas there was no significant activation in this region when semantically anomalous sentences were presented. Moreover, identical stimuli of normal sentences elicited activation in the left inferior frontal gyrus, only when the employment of syntactic knowledge was required. This task‐selective activation was not observed in any other measured regions, including the right homologous region. These results demonstrate that OT techniques, when coupled with the event‐related paradigm, are useful for studying the higher cognitive functions of the human cerebral cortex. Hum. Brain Mapping 17:89–99, 2002.