Multiple penetrations in the somatosensory cortex of three anesthetized raccoons 1 week following amputation of the fourth digit provided detailed information about somatotopy and neuronal responsiveness in the deafferented cortex. Recordings in a total of 601 penetrations (292 in deafferented cortex and 309 in the surrounding cortex) were compared with those from intact control animals described previously (Rasmusson et al., 1991). The level of spontaneous activity increased within the deafferented cortex, with 42% of the sites having high or moderate levels of spontaneous activity, in comparison with 18% in control animals. There was also an increase in the incidence of inhibitory responses to stimulation of adjacent digits (26% of the penetrations vs. 10% in control animals), confirming previous findings. These two variables, increased spontaneous activity and the presence of strong lateral inhibition, were highly correlated in individual penetrations. An unexpected finding was that the cortex representing the intact parts of forepaw was also disrupted with respect to these two measures, suggesting that amputation had an effect outside the deafferented region. In contrast, response properties that are more clearly a reflection of information processing in the dorsal column-medial lemniscal pathway (adaptation and threshold) were altered only within the deafferented region. The deafferented region was not homogeneous immediately after amputation, but consisted of a radically affected core region and a slightly affected fringe adjacent to the intact representations. This inhomogeneity had also been apparent with partial digit deafferentation, reported previously. The fringe, approximately 1 mm in width, may reflect overlapping projections from adjacent digits at one or more levels of the somatosensory pathway. Since the size of the fringe is similar to the maximum extent of reorganization found in other models of reorganization, the mechanisms of plasticity within this region may involve an unmasking of pre-existing synapses with slight modification in synaptic strength. However, the plasticity within the core region of the raccoon seen in these experiments, which may be 5 mm from nondeafferented cortex, requires more extensive changes, perhaps via polysynaptic pathways.
A retractable wire knife was employed to transect the reticular formation at the pontomedullary junction in order to assess the respective importance of pontine and medullary reticular neurons and their pathways in paradoxical sleep. Thirteen cats were implanted with a standard array of electrodes for polygraphic recording of sleep-wakefulness states during 3 days in baseline condition and during 21 days after transections. Average electroencephalographic (EEG) amplitude, average electromyographic (EMG) amplitude, and ponto-geniculo-occipital (PGO) spike rate were measured per 1-min epoch for each day. A trivariate computer graphics display of 1 day's data revealed three major clusters of points that corresponded to wakefulness, slow wave sleep, and paradoxical sleep in baseline. (a) After transections through the entire reticular formation at the pontomedullary junction, paradoxical sleep was no longer evident in the trivariate computer graphics or polygraphic record, either by the presence of a high PGO spike rate or by that of muscle atonia in association with a low-amplitude EEG. These results indicated that the reticular fibers that pass through the pontomedullary junction and interconnect the pontine tegmentum and the medullary reticular formation are necessary for generating the cluster of electrographic variables that normally characterizes paradoxical sleep. (b) After transections through the dorsal half of the reticular formation, paradoxical sleep was still evident, though with a reduced PGO spike rate, and muscle atonia was normal. These results indicated that the descending noradrenaline locus coeruleus fibers and the "longitudinal catecholamine bundle," which course through the dorsal tegmentum, are not necessary for the generation of muscle atonia or the state of paradoxical sleep. (c) After transections through the ventral half of the reticular formation, paradoxical sleep was still apparent by the association of a moderate, though reduced, rate of PGO spiking in association with low-amplitude EEG activity and a high-amplitude EMG, indicating a loss of muscle atonia. The duration of the PS episodes, however, was greatly reduced. These results indicated that the descending "tegmentoreticular" and ascending reticulotegmental pathways, which course ventrally through the pontomedullary junction and interconnect the dorsolateral pontine tegmentum and the ventromedial medullary reticular formation, are essential for the muscle atonia of paradoxical sleep and important for the normal cyclic generation and maintenance of the state of paradoxical sleep.
Electrophysiological recordings were made at a large number of sites in the primary somatosensory cortex of six anesthetized raccoons. A high density of penetrations (110-229 per animal), within or near the representation of the fourth digit, allowed identification of three cortical regions with different physiological properties: a glabrous zone, containing a highly detailed, somatotopically ordered representation of the glabrous surface of the digit; rostral to this a claw-dominant zone, in which the neurons at most penetrations respond to stimulation of the claw of the fourth digit, but may also receive input from the hairy skin or surrounding glabrous skin; and a more rostral multidigit zone, in which the neurons respond to stimulation of two to five digits, with the dominant digit usually being the one represented caudally (i.e., the fourth digit at most of the sites sampled here). Claw-dominant zones with receptive fields restricted to digit three or five are also found rostral to the representations of the glabrous skin of the corresponding digit. The glabrous and claw-dominant zones constitute a complete map of the fourth digit. The multidigit region presumably is a separate map, since its neurons have different spatial convergence, higher thresholds, and a lower incidence of slowly adapting inputs than those in the claw-dominant and glabrous zones. A comparison between animals with lesions of the basal forebrain and intact animals found no differences in the organization of these zones or in the responses to peripheral input, suggesting that cholinergic inputs to the cortex are not essential to these properties. The detailed description of these regions and the proposed terminology should resolve some inconsistencies in the use of the term "heterogeneous zone" in this species.
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