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Somatosensory evoked potentials (SEPs) are generated in afferent pathways, subcortical structures and various regions of cerebellar and cerebral cortex by stimulation of somatic receptors or electrical stimulation of peripheral nerves. This review summarizes current knowledge of SEPs generated in cerebral cortex by stimulation of the median nerve, the most common form of stimulation for human research and clinical investigations. Major sources of data for the review are intracranial recordings obtained from patients during diagnostic or neurosurgical procedures, and similar recordings in monkeys. Short-latency cortical SEPs in the 20-40 ms latency range consist of P20 and N30, recorded from motor cortex and frontal scalp; P25 and N35, recorded from cortex near the central sulcus and central scalp; and N20 and P30, recorded from somatosensory cortex and parietal scalp. Several lines of evidence including cortical surface and intracerebral recordings, neuromagnetic recordings and lesion studies in humans and monkeys, strongly support the conclusion that these potentials are generated in contralateral somatosensory cortex in areas 3b and 1, in contrast to the conclusion of many previous studies that SEPs recorded from the frontal scalp are generated in motor cortex and other frontal lobe areas. These potentials are primarily mediated by cutaneous afferents of the dorsal column-medial lemniscal system; the contribution of muscle afferents has not been completely resolved but appears to be small. There is currently no evidence that short-latency SEPs are generated in cortex other than primary somatosensory cortex. Recordings from the vicinity of the second somatosensory area, from the supplementary motor and sensory areas and from surface cortex other than sensorimotor cortex have not detected reliable short-latency activity, although some of these regions generate long-latency potentials. Consequently, short-latency SEPs recorded from the scalp are similar to those recorded from the surface of sensorimotor cortex. Old World monkeys such as Macaca mulatta and M. fascicularis provide an excellent model for human short-latency SEPs. All the potentials described above in humans have corresponding monkey analogues, with similar distributions over the cortical surface. The squirrel monkey, a New World species, exhibits the same potentials, but due to the different morphology of sensorimotor cortex, the surface distribution of SEPs is quite different.

1. The anatomic generators of human median nerve somatosensory evoked potentials (SEPs) in the 40 to 250-ms latency range were investigated in 54 patients by means of cortical-surface and transcortical recordings obtained during neurosurgery. 2. Contralateral stimulation evoked three groups of SEPs recorded from the hand representation area of sensorimotor cortex: P45-N80-P180, recorded anterior to the central sulcus (CS) and maximal on the precentral gyrus; N45-P80-N180, recorded posterior to the CS and maximal on the postcentral gyrus; and P50-N90-P190, recorded near and on either side of the CS. 3. P45-N80-P180 inverted in polarity to N45-P80-N180 across the CS but was similar in polarity from the cortical surface and white matter in transcortical recordings. These spatial distributions were similar to those of the short-latency P20-N30 and N20-P30 potentials described in the preceding paper, suggesting that these long-latency potentials are generated in area 3b of somatosensory cortex. 4. P50-N90-P190 was largest over the anterior one-half of somatosensory cortex and did not show polarity inversion across the CS. This spatial distribution was similar to that of the short-latency P25-N35 potentials described in the preceding paper and, together with our and Goldring et al. 1970; Stohr and Goldring 1969 transcortical recordings, suggest that these long-latency potentials are generated in area 1 of somatosensory cortex. 5. SEPs of apparently local origin were recorded from several regions of sensorimotor cortex to stimulation of the ipsilateral median nerve. Surface and transcortical recordings suggest that the ipsilateral potentials are generated not in area 3b, but rather in other regions of sensorimotor cortex perhaps including areas 4, 1, 2, and 7. This spatial distribution suggests that the ipsilateral potentials are generated by transcallosal input from the contralateral hemisphere. 6. Recordings from the periSylvian region were characterized by P100 and N100, recorded above and below the Sylvian sulcus (SS) respectively. This distribution suggests a tangential generator located in the upper wall of the SS in the second somatosensory area (SII). In addition, N125 and P200, recorded near and on either side of the SS, suggest a radial generator in a portion of SII located in surface cortex above the SS. 7. In comparison with the short-latency SEPs described in the preceding paper, the long-latency potentials were more variable and were more affected by intraoperative conditions.(ABSTRACT TRUNCATED AT 400 WORDS)

1. The anatomic generators of human median nerve somatosensory evoked potentials (SEPs) in the 40 to 250-ms latency range were investigated in 54 patients by means of cortical-surface and transcortical recordings obtained during neurosurgery. 2. Contralateral stimulation evoked three groups of SEPs recorded from the hand representation area of sensorimotor cortex: P45-N80-P180, recorded anterior to the central sulcus (CS) and maximal on the precentral gyrus; N45-P80-N180, recorded posterior to the CS and maximal on the postcentral gyrus; and P50-N90-P190, recorded near and on either side of the CS. 3. P45-N80-P180 inverted in polarity to N45-P80-N180 across the CS but was similar in polarity from the cortical surface and white matter in transcortical recordings. These spatial distributions were similar to those of the short-latency P20-N30 and N20-P30 potentials described in the preceding paper, suggesting that these long-latency potentials are generated in area 3b of somatosensory cortex. 4. P50-N90-P190 was largest over the anterior one-half of somatosensory cortex and did not show polarity inversion across the CS. This spatial distribution was similar to that of the short-latency P25-N35 potentials described in the preceding paper and, together with our and Goldring et al. 1970; Stohr and Goldring 1969 transcortical recordings, suggest that these long-latency potentials are generated in area 1 of somatosensory cortex. 5. SEPs of apparently local origin were recorded from several regions of sensorimotor cortex to stimulation of the ipsilateral median nerve. Surface and transcortical recordings suggest that the ipsilateral potentials are generated not in area 3b, but rather in other regions of sensorimotor cortex perhaps including areas 4, 1, 2, and 7. This spatial distribution suggests that the ipsilateral potentials are generated by transcallosal input from the contralateral hemisphere. 6. Recordings from the periSylvian region were characterized by P100 and N100, recorded above and below the Sylvian sulcus (SS) respectively. This distribution suggests a tangential generator located in the upper wall of the SS in the second somatosensory area (SII). In addition, N125 and P200, recorded near and on either side of the SS, suggest a radial generator in a portion of SII located in surface cortex above the SS. 7. In comparison with the short-latency SEPs described in the preceding paper, the long-latency potentials were more variable and were more affected by intraoperative conditions.

The traditional means of localizing sensorimotor cortex during surgery is Penfield's procedure of mapping sensory and motor responses elicited by electrical stimulation of the cortical surface. This procedure can accurately localize sensorimotor cortex but is time-consuming and best carried out in awake, cooperative patients. An alternative localization procedure is presented that involves cortical surface recordings of somatosensory evoked potentials (SEP's), providing accurate and rapid localization in patients under either local or general anesthesia. The morphology and amplitude of median nerve SEP's recorded from the cortical surface varied systematically as a function of spatial location relative to the sensorimotor hand representation area. These results were validated in 18 patients operated on under local anesthesia in whom the sensorimotor cortex was independently localized by electrical stimulation mapping; the two procedures were in agreement in all cases. Similar SEP results were demonstrated in an additional 27 patients operated on under general anesthesia without electrical stimulation mapping. The following three spatial relationships between SEP's and the anatomy of the sensorimotor cortex permit rapid and accurate localization of the sensorimotor hand area: 1) SEP's with approximately mirror-image waveforms are recorded at electrode sites in the hand area on opposite sides of the central sulcus (P20-N30 precentrally and N20-P30 postcentrally); 2) the P25-N35 is recorded from the postcentral gyrus as well as a small region of the precentral gyrus in the immediate vicinity of the central sulcus: this waveform is largest on the postcentral gyrus about 1 cm medial to the focus of the 20- and 30-msec potentials; and 3) regardless of component identification, maximum SEP amplitudes are recorded from the hand representation area on the precentral and postcentral gyri.

Task-dependent field potentials were recorded from implanted electrodes located in the hippocampus and other medial temporal lobe (MTL) structures of epileptic patients undergoing evaluation for possible surgery. In 2-alternative categorization tasks, low-probability auditory, somatic, and visual stimuli elicited potentials with large amplitudes and sharp spatial gradients having the following characteristic spatial distribution: positive posterior to the hippocampus, negative within the hippocampus, and positive anterior to the hippocampus. The sharp spatial gradients within the MTL suggest that these potentials were locally generated, probably by hippocampal pyramidal cells. The MTL potentials were also reliably elicited by exemplars of semantic categories and by stimulus omissions and were sensitive to the sequence of preceding stimuli. However, they were not elicited by the same stimulus sequences when the patient's attention was directed elsewhere and categorization was not required. These results indicate that the MTL potentials reflect endogenous as opposed to obligatory processes. The time course and task dependence of the MTL potentials suggest that MTL structures could contribute to P300 and related event-related potentials on the scalp.

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