The combination of transcranial magnetic stimulation (TMS) with functional neuroimaging has expanded the potential of TMS for human brain mapping. The precise and reliable positioning of the TMS coil is not a simple task, however. Modern frameless stereotaxic systems allow investigators to base navigation either on the subject's structural magnetic resonance imaging (MRI), functional MRI data, or the use of functional neuroimaging data from the literature, so-called ''probabilistic approach.'' The latter assumes consistency across individuals in the location of task-related ''activations'' in standardized stereotaxic space. Conventional nonstereotaxic localization of brain areas is also a common method for defining the coil position. Our aim was to evaluate the accuracy of five different localization strategies in one single study. The left primary motor cortex (left M1-Hand) was used as target region. Three approaches were based on real-time frameless stereotaxy using information based on either anatomical or functional MRI. The remaining two strategies relied either on standard cranial landmarks (i.e., the International 10-20 EEG system) or a standardized function-guided procedure (i.e., the spatial relationship between the left and right M1-Hand). The results were compared to a TMS-based mapping of the primary motor cortex; center of gravity of motor-evoked potentials (MEP-CoG) was calculated for each subject (n ¼ 10). Our findings suggest that highest precision can be achieved with fMRI-guided stimulation, which was accurate within the range of millimeters. Very consistent results were also obtained with the ''probabilistic'' approach. In view of these findings, we discuss the methods and special characteristics of each localization strategy.

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Human speech and hand use both involve highly specialized complex movement patterns. Whereas previous studies in detail characterized the cortical motor systems mediating speech and finger movements, the network that provides coordination of concurrent speech and hand movements so far is unknown. Using functional magnetic resonance imaging (fMRI), the present study investigated differential cortical networks devoted to speech or fingertapping, and regions mediating integration of these complex movement patterns involving different effectors. The conjunction contrasts revealing regions activated both during sole fingertapping and sole repetitive articulation or reading aloud showed contralateral regions at the border of ventral and dorsal motor cortex. In contrast, the analyses revealing regions showing a higher level of fMRI activation for concurrent movements of both effectors compared with sole hand movements or repetitive articulation or reading aloud showed distinct premotor activations, which were situated dorsal and caudal to the areas activated across speech and fingertapping tasks. These results indicate that the premotor cortex (PMC) subserves coordination of concurrent speech with hand movements. This integrative motor region is not identical with the area that shows overlapping activations for speech and fingertapping. Thus, concurrent performance of these complex movement patterns involving different effectors requires, in addition to somatotopic motor cortex activation, orchestration subserved by a distinct PMC area.

Sparing R, Dafotakis M, Buelte D, Meister IG, Noth J. Excitability of human motor and visual cortex before, during, and after hyperventilation. J Appl Physiol 102: 406 -411, 2007. First published September 21, 2006; doi:10.1152/japplphysiol.00770.2006.-In humans, hyperventilation (HV) has various effects on systemic physiology and, in particular, on neuronal excitability and synaptic transmission. However, it is far from clear how the effects of HV are mediated at the cortical level. In this study we investigated the effects of HV-induced hypocapnia on primary motor (M1) and visual cortex (V1) excitability. We used 1) motor threshold (MT) and phosphene threshold (PT) and 2) stimulus-response (S-R) curves (i.e., recruitment curves) as measures of excitability. In the motor cortex, we additionally investigated 3) the intrinsic inhibitory and facilitatory neuronal circuits using a short-interval paired-pulse paradigm. Measurements were performed before, during, and after 10 min of HV (resulting in a minimum end-tidal PCO2 of 15 Torr). HV significantly increased motor-evoked potential (MEP) amplitudes, particularly at lower transcranial magnetic stimulation (TMS) intensities. Pairedpulse stimulation indicated that HV decreases intracortical inhibition (ICI) without changing intracortical facilitation. The results suggest that low PCO 2 levels modulate, in particular, the intrinsic neuronal circuits of ICI, which are largely mediated by neurons containing ␥-aminobutyric acid. Modulation of MT probably resulted from alterations of Na ϩ channel conductances. A significant decrease of PT, together with higher intensity of phosphenes at low stimulus intensities, furthermore suggested that HV acts on the excitability of M1 and V1 in a comparable fashion. This finding implies that HV also affects other brain structures besides the corticospinal motor system. The further exploration of these physiological mechanisms may contribute to the understanding of the various HV-related clinical phenomenona. partial pressure of carbon dioxide; phosphenes; threshold; pairedpulse transcranial magnetic stimulation; intracortical facilitation; intracortical inhibition HYPERVENTILATION (HV) (or hyperpnea) is the state of breathing faster or deeper than necessary, thereby reducing the CO 2 concentration of the blood below normal. What is usually referred to as HV is, in fact, hypocapnia. Since a reduction of arterial PCO 2 (Pa CO 2 ) below the normal level (40 Torr) is obtained by increasing the alveolar ventilation, HV became synonymous with hypocapnia (32). HV is known to have various effects on human physiology (for a review, see Ref. 6).For instance, a reduction in Pa CO 2 increases the excitability of sensory and motor axons in the peripheral nervous system (21,23,28).The aim of the present study was to investigate further the neural mechanisms of the HV-induced changes in cortical excitability. As measures of primary motor cortex (M1) excitability, motor threshold (MT), stimulus-response (S-R) curves (i.e., recruitment curves), intra...

Although language functions are, in general, attributed to the left hemisphere, it is still a matter of debate to what extent the cognitive functions underlying the processing of music are lateralized in the human brain. To investigate hemispheric specialization we evaluated the effect of different overt musical and linguistic tasks on the excitability of both left and right hand motor cortices using transcranial magnetic stimulation (TMS). Task-dependent changes of the size of the TMS-elicited motor evoked potentials were recorded in 12 right-handed, musically naive subjects during and after overt speech, singing and humming, i.e. the production of melody without word articulation. The articulation of meaningless syllables served as control condition. We found reciprocal lateralized effects of overt speech and musical tasks on motor cortex excitability. During overt speech, the corticospinal projection of the left (i.e. dominant) hemisphere to the right hand was facilitated. In contrast, excitability of the right motor cortex increased during both overt singing and humming, whereas no effect was observed on the left hemisphere. Although the traditional concept of hemispheric lateralization of music has been challenged by recent neuroimaging studies, our findings demonstrate that right-hemisphere preponderance of music is nevertheless present. We discuss our results in terms of the recent concepts on evolution of language and gesture, which hypothesize that cerebral networks mediating hand movement and those subserving language processing are functionally linked. TMS may constitute a useful tool to further investigate the relationship between cortical representations of motor functions, music and language using comparative approaches.