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com/locate/ynimg NeuroImage 25 (2005) 727 733

Cortical representation of dermatomes: MEG-derived maps after tactile stimulation

Eduardo M. Castillo* and Andrew C. Papanicolaou
Department of Neurosurgery, Vivian L. Smith Center for Neurologic Research, University of Texas Houston, Medical School, 1333 Moursund Street, Ste. H114, Houston, TX 77030, USA Received 19 April 2004; revised 14 December 2004; accepted 16 December 2004 Available online 16 February 2005 Mechanical stimulation of skin receptors is known to evoke cortical responses arising from the somatosensory cortex. Here we present a magnetoencephalographic (MEG) study where dermatomal somatosensory-evoked fields (DSSEFs) were recorded after mechanical stimulation of sacral (S1), lumbar (L3), thoracic (Th7), and cervical (C4) dermatomes in three healthy volunteers. All MEG measurements were repeated in order to test the replicability of the results. DSSEFs were successfully measured and modeled in all three participants. The topography and temporal dynamics of cortical responses derived after stimulation of each dermatome are described. We found that corticalevoked responses can be reliably recorded using MEG after mechanical stimulation of dermatomes when a sufficiently large skin region within the dermatome is stimulated. Primary sensory cortex response (SI) to each of the four dermatomes was replicable and showed stability over time. The MEG-derived individual maps of activation confirm the somatotopic representation of dermatomes in primary sensory cortex and the utility of MEG recordings in disentangling the interactions between primary and secondary sensory cortex during somatic perception. D 2005 Elsevier Inc. All rights reserved. Keywords: Dermatomes; Magnetoencephalography; Tactile stimulation

Introduction The sensations of touch, pressure, and vibration result from stimulation of tactile receptors in the skin. Impulses from those receptors first enter the peripheral afferent nerve fiber, then the spinal ganglion, then the dorsal roots, then the spinal cord, and finally the brain. After the peripheral nervous system is completely developed, each spinal nerve (delivered from a specific segment of the spinal cord) will innervate a well-defined area of the skin called

* Corresponding author. Fax: +1 713 7977590. E-mail address: eduardo.m.castillo@uth.tmc.edu (E.M. Castillo). Available online on ScienceDirect (www.sciencedirect.com). 1053-8119/$ - see front matter D 2005 Elsevier Inc. All rights reserved. doi:10.1016/j.neuroimage.2004.12.040

a dermatome. Thus, each dermatome represents a specific area of nerve reception of sensory impulses. It is well documented how cortical responses can be recorded after peripheral stimulation of the skin receptors (Chiappa and Cros, 1997; Hari and Forss, 1999; Kakigi et al., 2000). Different studies have focused on the description of dermatomal somatosensory-evoked potentials (DSSEPs) and the development of a consistent methodology to evoke and record these responses for the purpose of using them diagnostically (Liguori and Taher, 1991). Katifi and Sedgwick (1986) described and validated electrophysiological techniques for recording DSSEPs following the stimulation of the L5 and S1 dermatomes in a group of 54 subjects. These normative data have been extremely valuable when DSSEPs are used to investigate spinal cord pathologies. The segmental arrangement of dermatomes has been used as a map to determine the sacral (S), lumbar (L), thoracic (T), or cervical (C) level of the spinal lesion by the sensory loss (Curt and Dietz, 1996). Hyperesthesias and anesthesias confined to specific dermatomes are often symptoms of dorsal root lesions but can also be due to brain tumors affecting sensory cortices. Direct stimulation of the roots (by needle electrodes or superficial electrical stimulation) can be used to study root lesions (Nemecek et al., 2003; Storm and Kraft, 2004; Synek, 1986). Measurements of latency and amplitude of segmental and dermatomal SSEPs have been used in the diagnosis of lumbosacral radiculopathy (Dumitru et al., 1993; Rodriquez et al., 1987) and recent studies suggest that those recordings can be a valuable index in demonstrating the efficacy of lumbar root decompression therapy (Naguszewski et al., 2001). Interestingly, little attention has been paid to the description of the cortical topography of these DSSEPs. The first description of the actual cortical representation of dermatomes in the primary sensory cortex (SI) comes from the pioneering recordings of Penfield and colleagues, where the somatotopic representation of the human body was first drawn using electrocortical stimulation mapping procedures during brain surgery (Penfield and Boldrey, 1937; Penfield and Rasmussen, 1957). Although these findings have been verified and extended in new studies using similar invasive neurophysiological recordings (Baumgartner et al., 1991;

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Woolsey et al., 1979), few complete references can be found describing the somatotopic arrangement of dermatomes. Novel neuroimaging techniques such as FMRI, PET, and MEG provide a unique opportunity to characterize the cortical organization of sensory responses. MEG has undergone a rapid evolution during the past two decades and has emerged, especially when high temporal resolution is desired, as the non-invasive method of choice for mapping the primary sensory cortex. Individual MEG-derived maps of activation have achieved an adequate agreement with invasive bgold standardQ procedures to delineate presurgically the representation of sensorimotor functions (Castillo et al., 2004; Ganslandt et al., 1999; Morioka et al., 1995; Rezai et al., 1997). The accuracy of the technique for localizing the central sulcus as compared with intraoperative electrocorticography is of the order of a few millimeters. These factors, coupled with the possibility to replicate (as many times as necessary) the measurement, renders MEG the tool of preference to study and disentangle the cortical representation of dermatomes in the human brain. Many studies using MEG have verified the somatotopic arrangement of body representation in the primary sensory cortex after stimulation of fingers (Hari et al., 1990; Pollok et al., 2002), lips (Disbrow et al., 2003), toes (Nakamura et al., 1998), tongue (Karhu et al., 1991), mouth (Disbrow et al., 2003), and ears (Nihashi et al., 2001), with little attention paid to the representation of the major thoracic or lumbar dermatomes. Although it is well documented how neurological conditions (especially tumors and AVMs) can produce sensory deficits and alter the somatosensory homunculus (Duffau et al., 2003; Papanicolaou et al., 2001; Roux et al., 1998; Vates et al., 2002), a replicable procedure to obtain and localize DSSEFs after mechanical stimulation has not yet been described. Up to date, attempts to record and localize physiological responses after mechanical stimulation of dermatomes have been especially discouraging. This fact may be due to technical difficulties in the recording and/or analysis processes. The poor signal to noise ratio (S/N) of recordings when tactile stimulation is applied to the dermatomes has been identified as the main limiting factor by some investigators (Nakamura et al., 1998). Cortical responses to air-puff stimulation are weaker than responses elicited by electrical stimulation (Forss et al., 1994). Therefore, most

studies of the cortical representation of dermatomes have been limited to recordings of responses to electrical stimulation (Itomi et al., 2000; Liguori and Taher, 1991). This approach maximizes the S/N but reduces the actual contribution of pressure receptors (i.e., mechanoreceptors) to the derived sensory input and subsequent cortical maps. Here we present the spatiotemporal maps of cortical activation derived after tactile stimulation of four dermatomal sites (S1: posterior calf; L3: anterior thigh; Th7: anterior thorax; and C4: anterior shoulder) in three healthy volunteers. To our knowledge, this study is the first to replicate the somatotopy of representation of dermatomes combining tactile stimulation and MEG recordings. We bypassed the main methodological problem, poor signal to noise ratio when such sites are stimulated, by using a wider stimulation area (within the limits of each dermatome) and a longer and variable interval of stimulation in order to avoid the habituation response of the skin receptors. Maps of activation were obtained using a 248 channel magnetoencephalography (MEG) and replicated in order to address the stability of the results. MEG has evolved in the last few years, providing highdensity whole-head systems able to record weaker signals from the brain structures. This feature provides the opportunity to characterize, in a non-invasive fashion, the topography of cortical responses evoked after tactile stimulation of dermatomes. We hope that this procedure will help to evaluate the integrity of afferent tactile input fibers in patients with spinal nerve and root lesions and open a new window to the study of batypicalQ cortical representation of dermatomes in neurological disorders.

Materials and method Participants One right-handed male (28 years old) and two females (33 and 36 years of age), one right-handed and one left-handed, volunteered to serve as participants. None of them had a history of neurological problems. The procedures used in the study were approved by The University of Texas-Houston Institutional

Fig. 1. Evoked magnetic responses recorded after stimulation of dermatome L3 (left anterior thigh) in participant 1. The field distribution (right) of the earliest component (peaking at ~50ms in this case) showed a clear dipolar distribution over the right central area.

E.M. Castillo, A.C. Papanicolaou / NeuroImage 25 (2005) 727733 Table 1 Mean (and standard deviation) of the latency (ms) and amplitude (RMS: root mean square; and Q: dipole strength) values of the initial component of the dermatomal responses (S1, L3, Th7, C4) for our three participants over the two replications Latency C4 Th7 L3 S1 25.6 35.7 48.8 61.4 (3.2) (3.9) (6.3) (11.5) RMS 36.2 19.0 23.4 21.4 (7.2) (6.2) (4.4) (11.6) Q 11.9 (2.9) 6.7 (4.3) 10.2 (5.2) 7.2 (3.7)

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Table 2 Estimates location (x , y, and z coordinates) of ECDs of SI response after stimulation of S1, L3, Th7, and C4 dermatomes in each participant S1 Subject 1 x 1.4 (0.1) y 0.6 (0) z 8.6 (0.2) Subject 2 x 1.7 (0.1) y 0.4 (0.3) z 8.8 (0.1) Subject 3 x 1.3 (0.3) y 0.3 (0.1) z 8.4 (0.1) L3 1.4 (0.3) 0.3 (0.1) 9.2 (0.1) 1.5 (0.3) 0.45 (0.1) 9.25 (0.1) 1.5 (0.1) 0.4 (0.3) 9.0 (0.2) Th7 1.7 (0.1) 0.8 (0) 9.4 (0.1) 2.1 (0.3) 0.9 (0.1) 9.15 (0.2) 1.7 (0.4) 0.9 (0.1) 8.8 (0.1) C4 2.5 (0.6) 1.8 (0.1) 8.8 (0.1) 2.4 (0.2) 1.6 (0) 9.2 (0.1) 2.4 (0.4) 1.3 (0.2) 8.7 (0.1)

This initial component was always located in the primary somatosensory cortex contralateral to the stimulation site.

Review Board and all participants signed a consent form after the nature of the procedures had been explained to them. Materials

Stimuli To ensure that a sufficient number of skin receptors were stimulated, a special pneumatic air stimulator (12 cm in diameter) was made ad hoc in order to cover enough area within each dermatome. This stimulator was covered by a flexible membrane (latex) allowing the delivery of uniform pressure over the skin surface during each stimulation. We stimulated four sites (i.e., dermatomes) on the left side of the body. The location of the four dermatomes (S1, L3, Th7 and C4) was based on the dermatome maps in the textbook by Daube and Sandok (1978). Stimulation was performed on the left side of the body. The stimulator was fixed to each stimulation point using surgical tape. Pneumatically driven mechanical taps (25 lb/in2) were applied to the dermatomes with a balloon diaphragm 12-cm in diameter. Stimulus duration was 40 ms (with a rise time of 20 ms); interstimulus interval was pseudo-randomized, varying from 0.7, 1.0, and 1.3 s. Stimuli were repeated 800 times for each of the four dermatomes. Patients were instructed to rest quietly during the recording sessions. They were told that a brief mechanical pressure pulse would be delivered by the pneumatic diaphragm placed on their bodies and that their task was to relax and remain still. A training block containing 10 stimuli was presented before the

Fig. 2. Estimated ECD of each dermatome (S1: green; L3: red; Th7: blue; and C4: yellow) derived from each of the two replications of the procedure (represented by dots of the same color) in our three participants. Stimulation was performed on the left side of the body and ECDs were found in the right postcentral gyrus (SI). There is a clear topographical arrangement of the dermatomal responses from infero-medial (S1) to more lateral (C4) positions.

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actual MEG response recording in order to ensure that the participant was able to perceive the stimulation in each dermatome. MEG recordings were repeated for each participant in order to test the replicability of the MEG measurements. MEG recordings and analysis MEG consists of recording changes in the magnetic flux timelocked to the onset of external stimuli. The principles of stimulation and MEG recording are the same as described for somatosensory mapping in previous studies (Papanicolaou, 1998; Papanicolaou et al., 2001). All MEG recordings were done using a whole-head neuromagnetometer containing an array of 248 gradiometers (4D 3600, San Diego). This system is situated in a sound-attenuating and magnetically shielded room. During MEG recordings participants were asked to keep their eyes open and avoid, as much as possible, blinking or otherwise moving their body. In all participants, data sets were acquired during stimulation of four dermatomes (S1, L3, Th7, and C4). Between 800 and 1000 epochs of 600 ms duration (200-ms prestimulus baseline) were acquired with a 1.0-Hz high-pass cutoff and a sampling rate of 508 Hz. DC was offset using the prestimulus period as a baseline. Epochs were time locked to stimulus onset, averaged (after visual inspection to reject those trials containing movement artifacts), and band-pass filtered (240 Hz) before additional analysis. The location, orientation, and amplitude of the best fitting single equivalent current dipole (ECD) were estimated using a spherical volume conductor model (Sarvas, 1987). Only activity sources accounting for N94% of the field variance and localized within volumes less than 4 cm3 were accepted. Magnetic resonance imaging (MRI) scans were obtained for all participants. Brain response sources represented as ECDs were superimposed on the individual T1-weighted MRIs to delineate the anatomical location. Anatomical landmarks (nasion and the two auricular points corresponding to the external meati) were localized in each individual and used for the co-registration of the MRI and MEG scans.

The amplitudes (root mean square, RMS) and latencies (at the peak of each component) of the evoked components were measured. The anatomical location of the corresponding dipole estimated for each component (measured at the peak) was defined after co-registration with structural MRI.

Results All three participants successfully underwent two MEG recording sessions (separated by at least 24 h) in sessions lasting 70 min or less. The morphology of the derived DSSEPs featured several components (see Fig. 1). The cortical topography of the resulting maps of activation (derived after dipole modeling of each component) involved ipsi- and contralateral sensory cortices. The first visible early component (i.e., deflection within the first 70 ms) was present in all participants after stimulation of all dermatomes and its source was always located in the postcentral gyrus (contralateral to the stimulation side). Since tactile information from skin mechanoreceptors is processed by areas 3b, 1, and 2 (Dykes et al., 1980), these areas could potentially be the neural generators of the recorded responses. The latencies of this early component show a clear relationship with the conduction distance as it is shown in Table 1. The topographic distribution of this early component (SI) for each dermatome is presented in Fig. 2. Sacral (S1), lumbar (L3), thoracic (Th7), and cervical (C4) dermatomes were distributed across the postcentral gyrus from ventral (S1) to more lateral (C4) sites in agreement with the previously described homunculus (Penfield and Boldrey, 1937). The procedure was repeated in all participants (on different days) to test the reproducibility of the waveforms and the resulting activation maps. Fig. 2 shows the replicability of the two ECDs computed at the peak of each SI component and for each of the four dermatomes. Table 2 shows the localization parameters (x , y, and z ) of the estimated ECDs for the earliest cortical response. Maximum ECD displacement across sessions ranged from 2 to 8 mm for this early response.

Table 3 Mean latencies (and standard deviations) and anatomical topography of the ECDs estimated for the evoked responses following dermatome stimulation in the three participants Participant 1 Dermatome C4 Th7 L3 S1 2 C4 Th7 L3 S1 3 C4 Th7 L3 S1 SI-c 21.5 (3.1), 97.2 (5.8) 33.6 (5.8) 51.9 (3.9) 56.2 (7.1), 72.4 (9.5) 28.9 78.0 34.6 49.3 (2.5), 58.4 (4.2), (8.1), 89.8 (4.3) (3.7), 97.0 (6.1) (5.1) SII-c 66.3 (6.2) 87.2 (7.9) 95.2 (2.1), 132.7 (11.3) 107.5 (11.3) 66.2 (3.9), 93.2 (4.8) 47.4 (3.7), 77.2 (7.4) 81.5 (5.9) SI-i 75.9 (10.4), 103.4 (21.1) 57.4 (3.5) 62.6 (4.4) 87.9 (7.8) 81.9 (9.2) 108.6 (9.4) SII-i 181.3 (6.6) 75.9 (8.1), 103.4 (13.1) 117.6 (11.5) 66.3 (8.3) 91.4 (7.8)

61.4 (4.3) 26.5 (3.4), 78.3 (9.3) 39.1 (2.1) 45.4 (5.1), 76.3 (5.3) 66.7 (5.9)

SI: primary somatosensory cortex; SII: secondary somatosensory cortex; c: contralateral; i: ipsilateral.

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Fig. 3. Averaged evoked fields (top), component-specific magnetic flux distributions (middle), and correspondent ECDs derived after stimulation of C4 in subject 2. Five clear components are identifiable in the averaged waveform (A, B, C, D, and E). The magnetic flux distribution at the peak of each component showed a clear dipolar morphology. ECDs calculated for each component showed the sequential flow of activation between contralateral primary sensory cortex (A, B, C, and D) and ipsilateral and contralateral secondary sensory cortex (C and E).

Patterns of activation of SI and SII areas Following the initial SI activation, a sequence of evoked field components and their sources were identified in SI and SII cortices. Repeated activation (i.e., when activity sources derived from independent component peaks were found in the same area) was a common finding in some of the areas (specifically in SI-c), as it was the engagement of ipsilateral and contralateral sensory cortices (SI and SII). Although the replicability of the measurements was excellent for each participant, not all participants showed the same profile of activation (i.e., sequence of cortical responses) following the initial contralateral SI activation. Table 3 summarizes the topography and mean latency values of those responses that were replicated (i.e., were present in the two recording sessions for each participant). In all cases, the localization of the ECDs derived after modeling the first visible component was found in contralateral parietal sensory cortices (SI), probably generated by proximal neuronal populations in areas 1, 3b, and 2. In some of the recordings, following contralateral SI activation (see Table 3), activation of secondary somatosensory cortex was present ipsilateral and contralateral to the stimulation (Fig. 3).

Discussion The main finding of this study is that DSSEFs can be recorded and modeled using MEG recordings after mechanical stimulation of skin receptors. To our knowledge, our study is the first to

delineate and replicate the topography of DSSEFs following mechanical stimulation. The MEG-derived maps of activation for the SI response verified to a great extent the proposed somatotopic arrangement of primary sensory cortex (i.e., homunculus). In agreement with previous studies (Itomi et al., 2000; Slimp et al., 1992), the latencies of the early dermatomal responses showed a clear relationship with the distances (and length of the conduction pathway) from the stimulus point to the cerebral cortex. Activation following the initial SI burst was found, engaging primary sensory cortices (ipsi and contralateral to the stimulation) and/or spreading to secondary somatosensory cortex (SII). The activation of SI and SII parietal areas during tactile perception has been previously reported and studied in detail using MEG (Hari and Forss, 1999; Hari et al., 1984, 1993; Hoechstetter et al., 2001). Similar patterns of activation have been described after stimulation of other body parts (Castillo et al., 2004; Hari et al., 1984) and may be part of the stimuli evaluation process (i.e., evaluation of the physical features of the stimuli) and sensorimotor integration (Huttunen et al., 1996). Individual differences in amplitude and presence of these responses can be related to neuroanatomical differences in the extension and localization of the receptive fields (Geyer et al., 2000), the experimental conditions, and/or cognitive factors affecting the stimuli evaluation (Papanicolaou et al., 1989). In our study, we were especially interested in developing a procedure able to characterize DSSEFs evoked after stimulation of pressure receptors since many neurological conditions can produce selective loss of tactile sensitivity. Previous attempts (some of them in our laboratory) to standardize a procedure to evoke and record DSSEFs after mechanical stimulation have been partially unsuc-

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E.M. Castillo, A.C. Papanicolaou / NeuroImage 25 (2005) 727733 Castillo, E.M., Simos, P.G., Wheless, J.W., Baumgartner, J.E., Breier, J.I., Billingsley, R.L., Sarkari, S., Fitzgerald, M.E., Papanicolaou, A.C., 2004. Integrating sensory and motor mapping in a comprehensive MEG protocol: clinical validity and replicability. NeuroImage 21, 973 983. Chiappa, K.H., Cros, D., 1997. Dermatomal somatosensory evoked potentials. In: Chiappa, K.H. (Ed.), Evoked Potentials in Clinical Medicine. Lippincott-Raven, Philadelphia, pp. 471 475. Curt, A., Dietz, V., 1996. Traumatic cervical spinal cord injury: relation between somatosensory evoked potentials, neurological deficit, and hand function. Arch. Phys. Med. Rehabil. 77, 48 53. Daube, J.R., Sandok, B.A., 1978. The Spinal Levels. Medical Neurosciences. Little Brown, Boston, pp. 296 322. 1987. Disbrow, E.A., Hinkley, L.B., Roberts, T.P., 2003. Ipsilateral representation of oral structures in human anterior parietal somatosensory cortex and integration of inputs across the midline. J. Comp. Neurol. 22, 487 495. Duffau, H., Capelle, L., Denvil, D., Sichez, N., Gatignol, P., Taillandier, L., Lopes, M., Mitchell, M.C., Roche, S., Muller, J.C., Bitar, A., Sichez, J.P., van Effenterre, R., 2003. Usefulness of intraoperative electrical subcortical mapping during surgery for low-grade gliomas located within eloquent brain regions: functional results in a consecutive series of 103 patients. J. Neurosurg. 98, 764 778. Dumitru, D., Newton, B.Y., Dreyfuss, P., 1993. Segmental v dermatomal somatosensory-evoked potentials. Normal intertrial variation and sideto-side comparison. Am. J. Phys. Med. Rehabil. 72, 75 83. Dykes, R.W., Rasmusson, D.D., Hoeltzell, P.B., 1980. Organization of primary somatosensory cortex in the cat. J. Neurophysiol. 43, 1527 1546. Forss, N., Salmelin, R., Hari, R., 1994. Comparison of somatosensory evoked fields to airpuff and electric stimuli. Electroencephalogr. Clin. Neurophysiol. 92, 510 517. Ganslandt, O., Fahlbusch, R., Nimsky, C., Kober, H., Moller, M., Steinmeier, R., Romstock, J., Vieth, J., 1999. Functional neuronavigation with magnetoencephalography: outcome in 50 patients with lesions around the motor cortex. J. Neurosurg. 91, 73 79. Geyer, S., Schormann, T., Mohlberg, H., Zilles, K., 2000. Areas 3a, 3b, and 1 of human primary somatosensory cortex. Part 2. Spatial normalization to standard anatomical space. NeuroImage 11, 684 696. Hari, R., Forss, N., 1999. Magnetoencephalography in the study of human somatosensory cortical processing. Philos. Trans. R. Soc. Lond., B Biol. Sci. 29, 1145 1154. Hari, R., Reinikainen, K., Kaukoranta, E., Hamalainen, M., Ilmoniemi, R., Penttinen, A., Salminen, J., Teszner, D., 1984. Somatosensory evoked cerebral magnetic fields from SI and SII in man. Electroencephalogr. Clin. Neurophysiol. 57, 254 263. Hari, R., Hamalainen, H., Hamalainen, M., Kekoni, J., Sams, M., Tiihonen, J., 1990. Separate finger representations at the human second somatosensory cortex. Neuroscience 37, 245 249. Hari, R., Karhu, J., Hamalainen, M., Knuutila, J., Salonen, O., Sams, M., Vilkman, V., 1993. Functional organization of the human first and second somatosensory cortices: a neuromagnetic study. Eur. J. Neurosci. 5, 724 734. Hoechstetter, K., Rupp, A., Stancak, A., Meinck, H.M., Stippich, C., Berg, P., Scherg, M., 2001. Interaction of tactile input in the human primary and secondary somatosensory cortexa magnetoencephalographic study. NeuroImage 14, 759 767. Huttunen, J., Wikstrom, H., Korvenoja, A., Seppalainen, A.M., Aronen, H., Ilmoniemi, R.J., 1996. Significance of the second somatosensory cortex in sensorimotor integration: enhancement of sensory responses during finger movements. NeuroReport 10, 1009 1012. Itomi, K., Kakigi, R., Maeda, K., Hoshiyama, M., 2000. Dermatome versus homunculus; detailed topography of the primary somatosensory cortex following trunk stimulation. Clin. Neurophysiol. 111, 405 412. Kakigi, R., Hoshiyama, M., Shimojo, M., Naka, D., Yamasaki, H., Watanabe, S., Xiang, J., Maeda, K., Lam, K., Itomi, K., Nakamura, A., 2000. The somatosensory evoked magnetic fields. Prog. Neurobiol. 61, 495 523.

cessful due to technical difficulties. Like other investigators (Nakamura et al., 1998), we found that MEG recordings following mechanical stimulation of dermatomes were almost flat due to the poor signal to noise ratio. In response to these findings, we speculated that the stimulation devices may have been inadequate to the type of receptors. We further theorized that the stimuli were too weak, or too focal and therefore, were not able to activate the receptors in the dermatome sufficiently. A continuous or repetitive stimulus causes the intensity of the subjective sensation to gradually decrease since the number of action potentials produced after the beginning of continual stimulation decreases (Marzi and Tassinari, 1984; Reznik, 1996). Eventually, pressure receptors do not respond at all, and the subsequent recordings of evoked responses are unsatisfactory. To test these assumptions, in this study we introduced a novel procedure that maximizes the signal to noise ratio and permits the recording and modeling of sensoryevoked responses derived after tactile stimulation of dermatomes. This improvement is based on the use of a skin stimulator that covers a wide area of each dermatome and delivers uniform pressure over the skin surface. As the strength and the area of the sensory stimulus increases, more receptors are activated, including those with higher thresholds. Consequently, more units fire simultaneously, more afferent pathways are activated, and this directly affects the strength of the signal arising from the primary somatosensory cortices. The procedure that we tested in our study allows clear and replicable recordings of DSSEFs without the need for electrical stimulation. Given that even those studies attempting the recording of DSSEFs after electrical stimulation have reported poor responses after stimulation of some of the dermatomes (Itomi et al., 2000), our procedure introduces a new method that maximizes the quality of recordings in a non-invasive fashion. Although in our study, like other investigators (Itomi et al., 2000; Nakamura et al., 1998), we followed anatomical landmarks described by Daube and Sandok (1978) to identify each dermatome, this approach does not guarantee that the stimulation was confined to a single dermatome and therefore should be improved upon. We conclude that the cortical representation of dermatomal areas can be addressed using MEG after mechanical stimulation with very precise temporal resolution and with adequate spatial precision. The presented data extend the results of previous studies using MEG recordings, clarifying the spatiotemporal course of brain activation after dermatomal stimulation and introducing a new procedure that improves non-invasive recordings of DSSEFs. Further investigations will be required to disentangle the functional interactions of SI and SII cortices during perception of dermatomal stimulation briefly described in our results. Acknowledgments This study was supported by the NIH Grant RO1N53794101A1 to Andrew C. Papanicolaou. We thank Dr. Mark Mc Manis and Shelly Fitzgerald for technical assistance and Hermann Hospital for the use of their facilities. References
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