Cortical and subcortical brain effects of transcranial magnetic stimulation (TMS)-induced movement: an interleaved TMS/functional magnetic resonance imaging study.

BACKGROUND: To date, interleaved transcranial magnetic stimulation and functional magnetic resonance imaging (TMS/fMRI) studies of motor activation have not recorded whole brain patterns. We hypothesized that TMS would activate known motor circuitry with some additional regions plus some areas dropping out. METHODS: We used interleaved TMS/fMRI (11 subjects, three scans each) to elucidate whole brain activation patterns from 1-Hz TMS over left primary motor cortex. RESULTS: Both TMS (110% motor threshold) and volitional movement of the same muscles excited by TMS caused blood oxygen level-dependent (BOLD) patterns encompassing known motor circuitry. Additional activation was observed bilaterally in superior temporal auditory areas. Decreases in BOLD signal with unexpected post-task "rebounds" were observed for both tasks in the right motor area, right superior parietal lobe, and in occipital regions. Paired t test of parametric contrast maps failed to detect significant differences between TMS- and volition-induced effects. Differences were detectable, however, in primary data time-intensity profiles. CONCLUSIONS: Using this interleaved TMS/fMRI technique, TMS over primary motor cortex produces a whole brain pattern of BOLD activation similar to known motor circuitry, without detectable differences from mimicked volitional movement. Some differences may exist between time courses of BOLD intensity during TMS circuit activation and volitional circuit activation.     

Corticospinal control of the thumb-index grip depends on precision of force control: a transcranial magnetic stimulation and functional magnetic resonance imagery study in humans

The corticospinal system (CS) is well known to be of major importance for controlling the thumb-index grip, in particular for force grading. However, for a given force level, the way in which the involvement of this system could vary with increasing demands on precise force control is not well-known. Using transcranial magnetic stimulation and functional magnetic resonance imagery, the present experiments investigated whether increasing the precision demands while keeping the averaged force level similar during an isometric dynamic low-force control task, involving the thumb-index grip, does affect the corticospinal excitability to the thumb-index muscles and the activation of the motor cortices, primary and non-primary (supplementary motor area, dorsal and ventral premotor and in the contralateral area), at the origin of the CS. With transcranial magnetic stimulation, we showed that, when precision demands increased, the CS excitability increased to either the first dorsal interosseus or the opponens pollicis, and never to both, for similar ongoing electromyographic activation patterns of these muscles. With functional magnetic resonance imagery, we demonstrated that, for the same averaged force level, the amplitude of blood oxygen level-dependent signal increased in relation to the precision demands in the hand area of the contralateral primary motor cortex in the contralateral supplementary motor area, ventral and dorsal premotor area. Together these results show that, during the course of force generation, the CS integrates online top-down information to precisely fit the motor output to the task's constraints and that its multiple cortical origins are involved in this process, with the ventral premotor area appearing to have a special role.     

Hemodynamic impairment as a stimulus for functional brain reorganization.

We used functional magnetic resonance imaging to investigate whether hemispheral hemodynamic impairment can play an independent role in the functional reorganization of motor-related activity in the brain. Fourteen patients with large vessel occlusion but no infarct performed a simple motor task with the hand contralateral to the occluded vessel. Statistical parametric maps of regional activity were generated to compare the distribution of motor-related activity among patients with that of control subjects. Patients were classified into normal or abnormal cerebral hemodynamics on the basis of intracerebral vasomotor reactivity using transcranial Doppler and carbon dioxide inhalation. Controls and patients with normal vasomotor reactivity showed typical motor activity in contralateral motor areas. When the 9 patients with abnormal vasomotor reactivity were compared with the 14 control subjects in a single analysis, unique motor activation was identified in ipsilateral motor regions in the nonhypoperfused hemisphere. In a confirmatory analysis, blood oxygen level-dependent (BOLD) signal intensity was averaged in prespecified motor regions of interest. A significant group by hemisphere interaction was identified, driven by higher ipsilateral and lower contralateral hemisphere BOLD signal in patients with abnormal vasomotor reactivity compared with controls (F=12.40, P=0.002). The average ipsilateral motor region signal intensity was also significantly higher in the subgroup of patients with abnormal vasoreactivity and no TIA compared with controls (P=0.04). Our results suggest that hemodynamic impairment in one hemisphere, even in the absence of any focal lesion or any symptoms can be associated with a functional reorganization to the opposite hemisphere.     

Exploring the after-effects of theta burst magnetic stimulation on the human motor cortex: a functional imaging study

Theta burst stimulation (TBS) is a protocol of subthreshold repetitive transcranial magnetic stimulation (rTMS) inducing changes in cortical excitability. From functional imaging studies with conventional subthreshold rTMS protocols, it remains unclear what type of modulation occurs (direction and dependency to neural activity) and whether putative effects are bound to unspecific changes in cerebral perfusion or require a functional challenge. In a within-subjects (n = 17) repeated measurement design including real TBS and a control session without stimulation, we examined neural activation in a choice-reaction task after application of intermittent TBS, a protocol, which enhances cortical excitability over the left motor cortex (M1). Brain activity was monitored by blood oxygenation level-dependent (BOLD) functional magnetic resonance imaging interleaved with measuring regional cerebral blood flow (rCBF) at rest using MR-based perfusion imaging. On a separate day, TMS-induced compound muscle action potentials (cMAPs) amplitude of the right hand was measured after excitatory TBS. Compared to control, a significant decrease in BOLD signal due to right hand motor activity during the choice-reaction task was observed mainly in the stimulated M1 and motor-related remote areas after stimulation. This decrease might represent a facilitating effect, because cMAPs amplitude increased upon TBS compared to control. No changes in rCBF at rest were observed. The data demonstrate that subthreshold intermittent TBS targets both the stimulated cortical area as well as remote areas. The facilitation changing the efficacy of neural signal transmission seems to be reflected by a BOLD signal decrease, whereas the network at rest does not appear to be affected.     

Eye movement control by the cerebral cortex

PURPOSE OF REVIEW: This review focuses on eye movement control by the cerebral cortex, mainly in humans. Data have emerged based on the important contribution of recent techniques such as transcranial magnetic stimulation and functional magnetic resonance imaging, which provide complementary results to those of the classical lesion and electrical stimulation studies. RECENT FINDINGS: The location of the human frontal eye field and its role in pursuit eye movement control were recently detailed. Cumulative evidence for the role of the dorsolateral prefrontal cortex in unwanted reflexive saccade inhibition, short-term spatial memory and prediction suggests that this area controls decisional processes governing ocular motor behaviour. The organization of spatial memory in the dorsolateral prefrontal cortex (short-term), the parahippocampal cortex (medium-term) and the hippocampal formation (long-term) is also reviewed with the results of recent transcranial magnetic stimulation studies. The relatively complicated anatomy of the posterior parietal cortex in humans is briefly described followed by some additional results concerning the location of the parietal eye field - within the posterior half of the intraparietal sulcus - and its role in visuo-spatial integration and attention. The other areas involved in spatial attention are also examined in the light of several recent contributing reports. Lastly, there are also new functional magnetic resonance imaging findings concerning the posterior cingulate cortex, which appears to be mainly involved in the control of externally guided eye movements and attentional mechanisms. SUMMARY: Many new findings on the organization of saccades and pursuit eye movements at the cortical level have recently been reported. Furthermore, eye movements are increasingly used as a tool to elucidate relatively complex neuropsychological processes such as attention, spatial memory, motivation and decisional processes, and a considerable number of reports dealing with these questions have been observed.     

Functional and biochemical alterations of the medial frontal cortex in obsessive-compulsive disorder

CONTEXT: The medial frontal cortex (MFC), including the dorsal anterior cingulate and the supplementary motor area, is critical for adaptive and inhibitory control of behavior. Abnormally high MFC activity has been a consistent finding in functional neuroimaging studies of obsessive-compulsive disorder (OCD). However, the precise regions and the neural alterations associated with this abnormality remain unclear. OBJECTIVE: To examine the functional and biochemical properties of the MFC in patients with OCD. DESIGN: Cross-sectional study combining volume-localized proton magnetic resonance spectroscopy and functional magnetic resonance imaging with a task encompassing inhibitory control processes (the Multi-Source Interference Task) designed to activate the MFC. SETTING: Healthy control participants and OCD patients recruited from the general community. PARTICIPANTS: Nineteen OCD patients (10 males and 9 females) and 19 age-, sex-, education-, and intelligence-matched control participants recruited from the general community. MAIN OUTCOME MEASURES: Psychometric measures of symptom severity, Multi-Source Interference Task behavioral performance, blood oxygen level-dependent activation, and proton magnetic resonance spectroscopy brain metabolite concentrations. RESULTS: Multi-Source Interference Task behavioral performance did not differ between OCD patients and control subjects. Reaction time interference and response errors were correlated with blood oxygen level-dependent activation in the dorsal anterior cingulate region in both groups. Compared with controls, OCD patients had greater relative activation of the supplementary motor area and deactivation of the rostral anterior cingulate during high- vs low-conflict (incongruent > congruent) trials. Patients with OCD also showed reduced levels of neuronal N-acetylaspartate in the dorsal anterior cingulate region, which was negatively correlated with their blood oxygen level-dependent activation of the region. CONCLUSIONS: Hyperactivation of the MFC during high- vs low-conflict conditions in patients with OCD may be a compensatory response to a neuronal abnormality in the region. This relationship may partly explain the nature of inhibitory control deficits that are frequently seen in this group and may serve as a focus of future treatment studies.     

名人面孔、动物和人造物类别特异性命名区脑定位的功能磁共振研究

目的研究名人面孔、动物和人造物3种不同类型命名任务的脑定位特点,为术中选择合适的命名任务提供依据。方法对21名健康志愿者行3种不同类型命名任务的血氧水平依赖功能磁共振(BOLD-fMRI)扫描,采用SPM8软件进行脑功能区活动图像分析,获取各激活区激活体素的位置和强度。结果名人面孔命名区主要位于双侧前颞叶包括海马和杏仁核,且存在左侧优势,同时还激活双侧颞下回三角部和盖部;命名动物主要激活左侧辅助运动区;而命名人造物主要激活左侧运动前区和右侧辅助运动区,3种任务均有双侧梭状回激活,但命名人造物激活区范围比其他2种任务都大。结论人、动物和人造物3种不同类型命名任务的脑定位存在差异,这提示在术中定位功能区过程中,需要选择不同类型的命名任务,以减少假阴性刺激,降低术后永久性功能障碍的发生率。     

Retinotopic mapping in the human visual cortex using vascular space occupancy-dependent functional magnetic resonance imaging

Recently, we introduced a new methodology, vascular space occupancy functional magnetic resonance imaging, which detects brain activation on the basis of blood volume changes in parenchymal microvasculature and may provide higher spatial specificity than the blood oxygenation level-dependent method. To study whether this technique can be used for advanced brain mapping applications, we performed retinotopic mapping using alternating horizontal and vertical wedges that stimulate different portions of the visual field. The results using vascular space occupancy functional magnetic resonance imaging showed clear boundaries for V1/V2/VP/V4v in the ventral areas and V1/V2/V3/V3A in the dorsal areas, similar to the maps obtained using blood oxygenation level-dependent functional magnetic resonance imaging. Vascular space occupancy functional magnetic resonance imaging is a useful addition to the other neuroimaging techniques. Disadvantages of vascular space occupancy functional magnetic resonance imaging include lower contrast-to-noise ratio (about 1/3 of that of blood oxygenation level-dependent method) and limited volume coverage (nine slices for TR=3 s).     

Functional magnetic resonance imaging and somatosensory evoked potentials in rats with a neonatally induced freeze lesion of the somatosensory cortex.

Brain plasticity is an important mechanism for functional recovery from a cerebral lesion. The authors aimed to visualize plasticity in adult rats with a neonatal freeze lesion in the somatosensory cortex using functional magnetic resonance imaging (fMRI), and hypothesized activation outside the primary projection area. A freeze lesion was induced in the right somatosensory cortex of newborn Wistar rats (n = 12). Sham-operated animals (n = 7) served as controls. After 6 or 7 months, a neurologic examination was followed by recording of somatosensory evoked potentials (SSEPs) and magnetic resonance experiments (anatomical images, fMRI with blood oxygen level-dependent contrast and perfusion-weighted imaging) with electrical forepaw stimulation under alpha-chloralose anesthesia. Lesioned animals had no obvious neurologic deficits. Anatomical magnetic resonance images showed a malformed cortex or hyperintense areas (cysts) in the lesioned hemisphere. SSEPs were distorted and smaller in amplitude, and fMRI activation was significantly weaker in the lesioned hemisphere. Only in a few animals were cortical areas outside the primary sensory cortex activated. The results are discussed in respect to an apparent absence of plasticity, loss of excitable tissue, the excitability of the lesioned hemisphere, altered connectivity, and a disturbed coupling of increased neuronal activity to the hemodynamic response.     

Functional magnetic resonance imaging in the assessment of motor centers in patients with brain tumors in motor area]

Magnetic resonance imaging (MRI) is become recognised as the most sensitive and specific imaging modality for the examination of central nervous system pathology. Blood oxygen level-dependent (BOLD) contrast imaging is a non-invasive functional MRI technique for localising active neuronal brain centres. The aim of our study was to determine usefulness of fMRI in detecting hand movements cortical activity in hemisphere with brain tumour and comparison with corresponding one. Six right-handed patients with brain tumours of central sulcus area, aged 20-50 years were examined using a commercial 1.5 T scanner. All patients underwent both conventional and functional magnetic resonance imaging (MRI) examinations. Simple hand movements were examined separately for right and left hand at a self-paced rate. Significant increase of signal intensity was found in: a) contralateral primary motor cortex in all cases during both motor tasks, b) ipsilateral primary motor cortex, supplementary motor cortex and premotor cortex of both hemispheres in a part of the cases c) displacement of the activity in the affected hemisphere in comparison to the opposite one was noticeable depending on the localisation and size of the tumour and accompanied oedema. Usefulness of functional MRI in detecting primary motor area in patients with brain tumours was proved. There is a difference between activation in affected cortex and corresponding normal cortex in the opposite hemisphere.     

In vivo measurement of blood oxygen saturation using magnetic resonance imaging: A direct validation of the blood oxygen level-dependent concept in functional brain imaging

A novel noninvasive magnetic resonance imaging (MRI) method was developed to determine in vivo blood oxygen saturation and its changes during motor cortex activation in small cerebral veins. Specifically, based on susceptibility measurements in the resting states, pial veins were found to have a mean oxygen saturation of Y_rest = 0.544 ± 0.029 averaged over 14 vessels in 5 volunteers. During activation, susceptibility measurements revealed an oxygen saturation change of ΔY_susc = 0.14 ± 0.02. Independent evaluation from blood flow velocity measurements yielded a value of ΔY_flow = 0.14 ± 0.04 for this change. These results validate the blood oxygenation level-dependent (BOLD) model in functional MRI (fMRI). Hum. Brain Mapping 5:341–346, 1997. © 1997 Wiley-Liss, Inc.     

Highly focal BOLD activation on functional MRI in a patient with progressive myoclonic epilepsy and diffuse giant somatosensory evoked potentials

We analyzed the effect of afferent input on patterns of brain electrical activation in a 31-year-old man with progressive myoclonic epilepsy (PME) by measuring the somatosensory evoked potential (SSEP) amplitude at the scalp after median nerve stimulation and examining the changes in the functional magnetic resonance imaging blood oxygen level-dependent (fMRI BOLD) signal. High-amplitude SSEPs were elicited at the wrist in association with highly focal BOLD activation of the contralateral sensorimotor areas. By contrast, no diffuse activation of either the frontal or the posterior parietal cortical areas was observed, as seen in previously recorded data on SSEPs from a healthy control group. The highly focal BOLD activation in this patient suggests that cortex hyperexcitability might be limited to the sensorimotor cortex in PME. The combined EEG-fMRI findings highlight a dissociation between BOLD activation and neurophysiological findings.     

Relation of Brain Stimulation Induced Changes in MEP Amplitude and BOLD Signal

BackgroundNon-invasive human brain stimulation can induce long-term plasticity reflected by changes in putative markers of synaptic activation, such as the motor evoked potential (MEP) amplitude elicited by transcranial magnetic stimulation or the task-dependent blood oxygenation level-dependent (BOLD) signal measured by functional magnetic resonance imaging.ObjectiveTo study the relationship between brain stimulation induced changes in MEP amplitude and BOLD signal.MethodsPaired associative stimulation of the hand area of the left primary somatosensory cortex (S1-PAS) was applied in 15 healthy subjects to induce excitability change in the adjacent primary motor cortex (M1) [Kriváneková et al. 2011, Eur J Neurosci 34:1292-1300]. Before and after S1-PAS, MEP amplitude in a right hand muscle, and the BOLD signal during a right hand motor or somatosensory activation task were measured.ResultsS1-PAS resulted in substantial individual MEP and BOLD signal changes, but these changes did not correlate in M1 or S1.ConclusionsFindings indicate that MEP amplitude and BOLD signal within the tested M1 reflect physiologically distinct aspects of synaptic excitability change. Therefore, it is suggested that MEP amplitude and BOLD signal are complementary rather than interchangeable markers of synaptic excitability.     

大脑中动脉梗死患者运动功能的恢复与脉络丛前动脉供血脑区有关？

no chinese abstract     

Recurrent ipsilateral hemiparesis in a patient with both uncrossed corticospinal tracts and reorganization of cortical motor areas – An opportune visitation of the motor tracts

We report the case of a patient who experienced recurrent ipsilateral hemiparesis in the setting of predominantly-uncrossed corticospinal tracts, with concomitant neuronal reorganization of the cortical motor maps, and the presence of aberrant interhemispheric connections. Their presence was supported by our results from diffusion tensor imaging tractography, functional magnetic resonance imaging, and transcranial magnetic stimulation. To our knowledge, this has never been reported before, and provides valuable insights into the mechanisms behind post-stroke motor recovery.     

An increased precision comparison of TMS-induced motor cortex BOLD fMRI response for image-guided versus function-guided coil placement.

OBJECTIVE: To examine with high precision the differences between function-guided and image-guided transcranial magnetic stimulation (TMS). METHOD: Using a calibrated TMS coil holder/positioner, interleaved TMS/functional magnetic resonance imaging (fMRI), and individualized anatomy-based regional normalization, we conducted a two-phase study of TMS coil positioning guided by either function (elicited thumb motion) or image-based targeting of the "hand knob," the anatomy associated with fMRI activation during thumb motion. RESULTS: In every case, image-guided TMS coil placement produced a thumb movement response at thresholds similar to those found under function guidance. Unexpectedly, function-guided coil locations clustered bimodally over central and precentral sulci. Image-guided locations clustered as anticipated toward the targeted gyral crown. Despite these differences, blood oxygenation level-dependent (BOLD) activation locations and magnitude for the two methods displayed no consistent differences in mean or variance between or within subjects. Image guidance produced more consistent coil placement from subject to subject relative to targeted anatomy. Surprisingly, BOLD time courses from image-guided experiments showed significantly slower return to baseline after TMS than was observed under function guidance. CONCLUSIONS: The results demonstrate the effectiveness and precision of image-guided positioning of TMS coils combined with a precisely adjustable holder/positioner and regional normalization. Image guidance provides an accurate TMS placement relative to individual anatomy when no external sign is available.     

Complex motor function in humans: validating and extending the postulates of Alexandr R. Luria.

OBJECTIVE: We used functional brain imaging to reevaluate Luria's postulates and to elaborate the neural circuitry underlying performance of complex motor tasks. BACKGROUND: The anatomic organization and physiologic functioning of the normal human motor system have great significance for understanding motor dysfunction and remediation in neurology. Working with victims of penetrating head injuries, noted Russian neuropsychologist Aleksandr R. Luria designed several tests of fine motor control to understand their difficulties with complex voluntary movements. This led to his postulates that such function involves the premotor cortices and their interaction with the parietal lobe. METHOD: Six healthy young adults performed the hand imitation, fist-scissors-gun, and piano key tasks during blood oxygen level-dependent functional magnetic resonance imaging at 3 T. RESULTS: All 3 tasks revealed activation of both premotor and parietal cortices. Furthermore, while hand Imitation relied more on the ventral premotor area and right parietal lobe, fist-scissors-gun and piano key relied more on the supplementary motor cortex. CONCLUSIONS: We postulate that differences in task-dependent activations across these tasks relate to degrees of sequential movement, pacing, and imitation. These results uphold Luria's original hypotheses, and extend that work by providing a further characterization of the motor areas involved in complex motor behaviors.     

Comparison of spatial and temporal pattern for fMRI obtained with BOLD and arterial spin labeling

Summary. Functional magnetic resonance imaging (fMRI) is presently either performed using blood oxygenation level-dependent (BOLD) contrast or using cerebral blood flow (CBF), measured with arterial spin labeling (ASL) technique. The present fMRI study aimed to provide practical hints to favour one method over the other. It involved three different acquisition methods during visual checkerboard stimulation on nine healthy subjects: 1) CBF contrast obtained from ASL, 2) BOLD contrast extracted from ASL and 3) BOLD contrast from Echo planar imaging. Previous findings were replicated; i) no differences between the three measurements were found in the location of the activated region; ii) differences were found in the temporal characteristics of the signals and iii) BOLD has significantly higher sensitivity than ASL perfusion. ASL fMRI was favoured when the investigation demands for perfusion and task related signal changes. BOLD fMRI is more suitable in conjunction with fast event related design.     

Neural correlates of antinociception in borderline personality disorder

CONTEXT: A characteristic feature of borderline personality disorder (BPD) is self-injurious behavior in conjunction with stress-induced reduction of pain perception. Reduced pain sensitivity has been experimentally confirmed in patients with BPD, but the neural correlates of antinociceptive mechanisms in BPD are unknown. We predicted that heat stimuli in patients with BPD would activate brain areas concerned with cognitive and emotional evaluation of pain. OBJECTIVE: To assess the psychophysical properties and neural correlates of altered pain processing in patients with BPD. DESIGN: Case-control study. SETTING: A university hospital. PARTICIPANTS: Twelve women with BPD and self-injurious behavior and 12 age-matched control subjects. INTERVENTIONS: Psychophysical assessment and blood oxygen level-dependent functional magnetic resonance imaging during heat stimulation with fixed-temperature heat stimuli and individual-temperature stimuli adjusted for equal subjective pain in all the participants. MAIN OUTCOME MEASURE: Blood oxygen level-dependent functional magnetic resonance imaging signal changes during heat pain stimulation. RESULTS: Patients with BPD had higher pain thresholds and smaller overall volumes of activity than controls in response to identical heat stimuli. When the stimulus temperature was individually adjusted for equal subjective pain level, overall volumes of activity were similar, although regional patterns differed significantly. Patient response was greater in the dorsolateral prefrontal cortex and smaller in the posterior parietal cortex. Pain also produced neural deactivation in the perigenual anterior cingulate gyrus and the amygdala in patients with BPD. CONCLUSION: The interaction between increased pain-induced response in the dorsolateral prefrontal cortex and deactivation in the anterior cingulate and the amygdala is associated with an antinociceptive mechanism in patients with BPD.     

Deep brain stimulation for Parkinson's disease dissociates mood and motor circuits: a functional MRI case study.

Behavioral disturbances have been reported with subthalamic (STN) deep brain stimulation (DBS) treatment in Parkinson's disease (PD). We report correlative functional imaging (fMRI) of mood and motor responses induced by successive right and left DBS. A 36-year-old woman with medically refractory PD and a history of clinically remitted depression underwent uncomplicated implantation of bilateral STN DBS. High-frequency stimulation of the left electrode improved motor symptoms. Unexpectedly, right DBS alone elicited several reproducible episodes of acute depressive dysphoria. Structural and functional magnetic resonance imaging (fMRI) imaging was carried out with sequential individual electrode stimulation. The electrode on the left was within the inferior STN, whereas the right electrode was marginally superior and lateral to the intended STN target within the Fields of Forel/zona incerta. fMRI image analysis (Analysis of Functional NeuroImages, AFNI) contrasting OFF versus ON stimulation identified significant lateralized blood oxygen level-dependent (BOLD) signal changes with DBS (P < 0.001). Left DBS primarily showed changes in motor regions: increases in premotor and motor cortex, ventrolateral thalamus, putamen, and cerebellum as well as decreases in sensorimotor/supplementary motor cortex. Right DBS showed similar but less extensive change in motor regions. More prominent were the unique increases in superior prefrontal cortex, anterior cingulate (Brodmann's area [BA] 24), anterior thalamus, caudate, and brainstem, and marked widespread decreases in medial prefrontal cortex (BA 9/10). The mood disturbance resolved spontaneously in 4 weeks despite identical stimulation parameters. Transient depressive mood induced by subcortical DBS stimulation was correlated with changes in mesolimbic cortical structures. This case provides new evidence supporting cortical segregation of motor and nonmotor cortico-basal ganglionic systems that may converge in close proximity at the level of the STN and the adjacent white matter tracts (Fields of Forel/zona incerta).