Progressive transneuronal changes in the brainstem and thalamus after long-term dorsal rhizotomies in adult macaque monkeys.

This study deals with a potential brainstem and thalamic substrate for the extensive reorganization of somatosensory cortical maps that occurs after chronic, large-scale loss of peripheral input. Transneuronal atrophy occurred in neurons of the dorsal column (DCN) and ventral posterior lateral thalamic (VPL) nuclei in monkeys subjected to cervical and upper thoracic dorsal rhizotomies for 13-21 years and that had shown extensive representational plasticity in somatosensory cortex and thalamus in other experiments. Volumes of DCN and VPL, number and sizes of neurons, and neuronal packing density were measured by unbiased stereological techniques. When compared with the opposite, unaffected, side, the ipsilateral cuneate nucleus (CN), external cuneate nucleus (ECN), and contralateral VPL showed reductions in volume: 44-51% in CN, 37-48% in ECN, and 32-38% in VPL. In the affected nuclei, neurons were progressively shrunken with increasing survival time, and their packing density increased, but there was relatively little loss of neurons (10-16%). There was evidence for loss of axons of atrophic CN cells in the medial lemniscus and in the thalamus, with accompanying severe disorganization of the parts of the ventral posterior nuclei representing the normally innervated face and the deafferented upper limb. Secondary transneuronal atrophy in VPL, associated with retraction of axons of CN neurons undergoing primary transneuronal atrophy, is likely to be associated with similar withdrawal of axons from the cerebral cortex and should be a powerful influence on reorganization of somatotopic maps in the somatosensory cortex.

Neurochemical organization of inferior pulvinar complex in squirrel monkeys and macaques revealed by acetylcholinesterase histochemistry, calbindin and Cat-301 immunostaining, and Wisteria floribunda agglutinin binding.

To investigate whether the inferior pulvinar complex has a common organization in different primates, the chemoarchitecture of the visual thalamus was re-examined in squirrel monkeys (Saimiri sciureus) and macaques (Macaca mulatta). The inferior pulvinar (PI) complex consisted of multiple subdivisions and encompassed the classic PI, and adjacent ventral parts of the lateral and medial pulvinar (PL and PM, respectively). In keeping with nomenclature suggested previously for macaques, the PI subdivisions were termed the posterior, medial, central, lateral, and lateral-shell (PI(P), PI(M), PI(C), PI(L), and PI(L-S)). In both species, PI(P) was intense for calbindin, light for acetylcholinesterase (AChE), and very light for Wisteria floribunda agglutinin (WFA) histochemistry. The PI(M) was calbindin poor, AChE rich, and moderate for WFA. The PI(C) was calbindin intense, lighter for AChE, and exhibited little WFA binding. PI(L) and PI(L-S) contained populations of large calbindin or WFA cells that were more numerous in PI(L-S). Although staining with the monoclonal antibody Cat-301 differed between macaques and squirrel monkeys, the same subdivisions were displayed. Moderately dense, patchy Cat-301 stain was found in PI(M) of macaques, whereas in squirrel monkeys PI(M) was light. Connections of the rostral dorsolateral (DLr) and middle temporal (MT) areas of visual cortex in squirrel monkeys were compared with PI subdivisions revealed by the newer histochemical methods in the same cases. The major connections of DLr were with PI(C) and of MT were with PI(M).

Human thalamus: neurochemical mapping of inferior pulvinar complex.

The primate pulvinar connects with the entire array of known visual areas and is postulated to play a role in selective visual attention. Recently, five separate neurochemical subdivisions of a region termed the inferior pulvinar (PI) complex were identified in monkeys. In the present study, similar histochemical procedures were applied to map the extent of the PI complex in humans. Acetylcholinesterase histochemistry and cytochrome oxidase staining demarcated four histochemical zones in human pulvinar, corresponding to the medial, central, lateral and lateral-shell (PI(M), PI(C), PI(L), and PI(L-S)) divisions of the PI complex in monkeys.

Subdivisions of the motor and somatosensory thalamus of primates revealed with Wisteria floribunda agglutinin histochemistry.

We obtained well-differentiated staining of thalamic subdivisions in rhesus macaques and squirrel monkeys using a lectin, Wisteria floribunda agglutinin (WFA), that labels extracellular matrix proteoglycans. Regional variations in staining were observed within the motor and somatosensory thalamic regions that bear on current interpretations of the organization of these regions. The pattern of WFA staining was generally similar to that obtained with Cat-301 antibody, which also stains proteoglycans. However, WFA reliably produced selective staining in both squirrel monkeys and macaques, whereas Cat-301 stained macaques more consistently than squirrel monkeys.

Area V1 in macaque monkeys projects to multiple histochemically defined subdivisions of the inferior pulvinar complex.

To investigate how visuotopic connections relate to chemoarchitecture of the inferior pulvinar (PI) complex in macaques, neuroanatomical tracers were placed into known portions of the visual representation in V1. Separate foci of label associated with both the upper and lower visual quadrants occupied neurochemically defined medial, central, lateral, and lateral-shell subdivisions, PIM, PIC, PIL, and PIL-S. Visuotopic connection patterns thus supported the concept of a larger PI that includes portions of three classically defined 'nuclei' [C. Gutierrez, A. Yaun and C.G. Cusick, Neurochemical subdivisions of the inferior pulvinar in macaque monkeys, J. Comp. Neurol., 363 (1995) 545-562.], and corresponds to the topographically organized V1 projection zone.

Overlapping and nonoverlapping cortical projections to cortex of the superior temporal sulcus in the rhesus monkey: double anterograde tracer studies.

To examine how fibers from functionally distinct cortical zones interrelate within their target areas of the superior temporal sulcus (STS) in the rhesus monkey, separate anterograde tracers were injected in two different regions of the same hemisphere known to project to the STS. Paired injections were placed in dorsal prearcuate cortex and the caudal inferior parietal lobule (IPL), interconnected regions that are part of a hypothesized distributed network concerned with visuospatial analysis or directed attention; in a presumed auditory region of the superior temporal gyrus (STG) and in extrastriate visual cortex, the caudal IPL and lower rim of the intraparietal sulcus; and in dorsal prearcuate cortex and the STG. Overlapping and nonoverlapping projections were then examined in STS visual and polysensory areas. Prefrontal and parietal fibers directly overlapped extensively in area MST and all subdivisions of presumed polysensory cortex (areas TPOc, TPOi, and TPOr), although nonoverlapping connections were also found. Although STG and IPL fibers targeted all TPO subdivisions, connections were to nonoverlapping, but often adjacent, columns. Paired prefrontal and STG injections revealed largely nonoverlapping vertical columns of connections but substantial overlap within layers VI and I or areas TPOc and TPOi. The findings suggest that area TPO contains differently connected modules that may maintain at least initial segregation of visual versus auditory inputs. Other modules within area TPO receive directly converging input from the posterior parietal and the prefrontal cortices and may participate in a distributed cortical network concerned with visuospatial functions.

Extensive cortical reorganization following sciatic nerve injury in adult rats versus restricted reorganization after neonatal injury: implications for spatial and temporal limits on somatosensory plasticity.

Expansion of the saphenous representation in rat S-I cortex following sciatic nerve injury, examined at different times after injury and following injury at different developmental stages, has contributed to the beginnings of a comprehensive view of spatial and temporal patterns of cortical reorganization. In the first few days or weeks after deafferentation in adult animals, cortical reorganization may be spatially constrained to convergence zones between central representations of peripheral nerves (Wall and Cusick, 1984; Garraghty et al., 1994a; Schroeder et al., 1995). The apparent steady state of the rat hindpaw system up to 5-6 months after sciatic nerve injury contrasts with the additional, nearly complete reorganization shown at times longer than 7-8 months. The late reorganization supports the concept that reorganized cortical maps can continue to be altered throughout life. The prolonged time course of change in the rat hindpaw system suggests that studies of "chronic" nerve injuries need to carefully define the reorganizational state of the system at the time intervals studied. For humans with peripheral nerve or amputation injuries, the results imply that the short term postinjury status can be further altered at longer times, perhaps decades later. To characterize the neurochemical consequences or mechanisms of cortical reorganization, it is necessary to consider possible differences between early versus late changes. Time dependent changes in neurotransmitters and their receptors have been described following peripheral injury (e.g., Avendaño et al., 1995). In addition, both early and late mechanisms or consequences of reorganization may differ spatially. In the example of changes in rat hindpaw cortex after sciatic nerve transection, neurochemical changes in "expansion" cortex may differ quantitatively or qualitatively from changes in the deafferented "sciatic dominant" zone. To accurately define neurochemical changes, it may thus be necessary to characterize sample zones as having intact or reorganized inputs, or as deprived of inputs. The studies of cortical reorganization following neonatal sciatic nerve injury underscore the importance of developmental age at time of injury. Most studies of critical periods in the central nervous system have emphasized greater plasticity in developing as opposed to adult animals. Early lesions or deprivation, however, not only result in connectional alterations, but can produce dramatically more atrophy or cell loss (e.g., see Cunningham, 1982; Waite, 1984; Himes and Tessler, 1989). A number of authors have commented on the seeming paradox of greater transneuronal and retrograde cell death, yet greater neuronal plasticity, in infant animals. How developmental stage influences plastic responses to peripheral injury in the somatosensory system is not completely understood. Early peripheral lesions may deprive central neurons of necessary trophic factors, accentuate naturally occurring central cell death, and thereby result in smaller central representations. Or, smaller central representations may be produced by competitive interactions of deprived with adjacent intact pathways. In addition, throughout all stages of development, the capacity for reorganization may be spatially limited and depend on the size or pattern of the peripheral injury.

Neurochemical subdivisions of the inferior pulvinar in macaque monkeys.

The architecture of the pulvinar of rhesus monkeys was investigated by acetylcholinesterase (AChE) histochemistry, and by immunocytochemistry for calbindin-D28k and the SMI-32 antibody. The presence of four inferior subdivisions, comparable to those found in architectonic-connectional studies in squirrel monkeys (C.G. Cusick, J.L. Scripter, J.G. Darensbourg, and J.T. Weber, 1993, J. Comp. Neurol. 336:1-30), provided a basis for a proposed revised terminology for visual sectors of the macaque pulvinar. In the present study, the inferior pulvinar (PI) was identified as a neurochemically distinct region that included the traditional cytoarchitectonic nucleus PI and adjacent portions of the lateral and medial pulvinar nuclei, PL and PM. In calbindin-D28k stains, the lateral subdivision of the inferior pulvinar (PIL) had less intense neuropil staining than the adjacent central division, PIC. The PIL was characterized by large, intensely immunopositive neurons seldom found within PIC. PIL occupied the traditional PL and PI and exhibited a narrow shell zone, PIL-S, restricted to PL. The medial division of the inferior pulvinar (PIM) was in a location previously shown to be strongly connected with the middle temporal visual area (MT) in macaques. PIM was found in the medial one-half of the traditional PI and extended into adjacent portions of the traditional PM and PL. PIM was distinguished by less intense neuropil staining for calbindin and many cells stained with the SMI-32 antibody for neurofilament protein. In AChE stains, PIL was moderately dark, PIC appeared lighter, and PIM was characterized by small, intensely stained patches. The small posterior division (PIP) stained darkly for calbindin, lightly for AChE, and was unstained with the SMI-32 antibody. Thus, neurochemical, and perhaps connectional, subdivisions exist within PI, the region of the pulvinar that relays information to striate, "lower order" extrastriate, and inferotemporal visual cortex.

Chemoarchitectonics and corticocortical terminations within the superior temporal sulcus of the rhesus monkey: evidence for subdivisions of superior temporal polysensory cortex.

Cortex of the upper bank of the superior temporal sulcus (STS) in macaque monkeys, termed the superior temporal polysensory (STP) region, corresponds largely to architectonic area TPO and is connectionally distinct from adjacent visual areas. To investigate whether or not the STP region contains separate subdivisions, immunostaining for parvalbumin and neurofilament protein (using the SMI-32 antibody) was compared with patterns of corticocortical terminations in the STS. Chemoarchitectonic results provided evidence for three caudal-to-rostral subdivisions: TPOc, TPOi, and TPOr. Area TPOc was characterized by patchy staining for parvalbumin and SMI-32 in cortical layers IV/III and III, respectively. Area TPOi had more uniform chemoarchitectonic staining, whereas area TPOr had a thicker layer IV than TPOi. The connectional results showed prefrontal cortex in the location of the frontal eye fields (area 8) and dorsal area 46 projected in a columnar pattern to all cortical layers of area TPOc, to layer IV of TPOi, and in a columnar fashion, with a moderate increase in density in layer IV, to TPOr. In TPOc, columns of frontal connections showed a periodicity similar to that of the SMI-32 staining. The caudal inferior parietal lobule (area 7a) and superior temporal gyrus projected to each subdivision of area TPO, displaying either panlaminar or fourth-layer terminations. In addition to STP cortex, parvalbumin and SMI-32 immunostaining allowed identification of caudal visual areas of the STS, including MT, MST, FST, and V4t. These areas received first- and sixth-layer projections from prefrontal cortex and area 7a.

Synaptic morphology of substance P terminals on catecholamine neurons in the commissural subnucleus of the nucleus tractus solitarii in the rat.

The ultrastructure of substance P-containing nerve terminals synapsing on catecholamine neurons in the rat commissural subnucleus of the nucleus tractus solitarii (NTScom) was studied using a double immunocytochemical labeling technique. Although there were numerous tyrosine hydroxylase-immunoreactive (TH-I) somata present, substance P immunoreactive (SP-I) cell bodies were only occasionally found in the NTScom. At the light microscopic level, many SP-I terminals were seen closely associated with TH-I dendrites and somata. At the electron microscopic level, SP-I terminals synapsing on TH-I structures were also readily encountered. SP-I terminals contained small, clear, and predominantly spherical vesicles (32 +/- 4 nm diameter), as well as large dense-cored vesicles approximately 100 nm in diameter. Postsynaptic TH-I dendritic profiles of various calibers and somata were encountered. These postsynaptic TH-I structures often showed postsynaptic densities. The morphological features of the SP-TH synapses in the present study, that is, the size of synaptic vesicles and the presence of postsynaptic densities, are quite different from those of central carotid sinus afferent synapses reported in our previous study [Chen et al. (1992), J. Neurocytol., 21:137-147]. Therefore, most of the SP terminals of the SP-TH synapses in the NTScom appear not to originate from the carotid sinus afferents. SP-I second-order neurons of the carotid sinus afferent pathway [Chen et al. (1991), J. Auton. Nerv. Syst., 33:97-98] may be one of the possible sources of such terminals.

Effects of chronic monocular enucleation on calcium binding proteins calbindin-D28k and parvalbumin in the lateral geniculate nucleus of adult rhesus monkeys.

The calcium binding proteins parvalbumin and calbindin-D28k were localized immunocytochemically within the lateral geniculate nucleus of adult monkeys at 1-7 months after monocular enucleation. Within the deafferented magno- and parvocellular layers, parvalbumin and calbindin-D28k immunoreactive fibers were depleted at all post-enucleation times. The neuronal staining for parvalbumin was similar in numerical density and intensity between the deafferented and intact layers. In hemispheres examined at 5 and 7 months post-enucleation, parvalbumin-immunoreactive fibers were also lost within the deprived ocular dominance bands in layers IVA, IVC and VI of the visual cortex, suggesting that cellular expression or axonal transport of parvalbumin may be decreased in the deafferented geniculate laminae. While the intact magno- and parvocellular layers contained very few neurons that were immunoreactive for calbindin-D28k, the density of calbindin-D28k-positive neurons increased in these layers after deafferentation. The counts of calbindin-D28k and parvalbumin immunostained neurons were not statistically different at 4-7 months post-enucleation. Because virtually all magno- and parvocellular projection neurons express parvalbumin, many parvalbumin neurons that normally do not contain calbindin-D28k may co-express this in response to injury. The findings suggest that long-term deafferentation imposes additional calcium buffering requirements on lateral geniculate neurons.

Chemoarchitectonic subdivisions of the visual pulvinar in monkeys and their connectional relations with the middle temporal and rostral dorsolateral visual areas, MT and DLr.

The organization of the inferior pulvinar complex (PI) in squirrel monkeys was studied with histochemical localization of the calcium binding proteins calbindin-D28k and parvalbumin, and of cytochrome oxidase. With each of these markers, the inferior pulvinar complex can be subdivided into four distinct regions. Calbindin-D28k immunoreactivity is densely distributed in cells and neuropil within PI, except for a distinct centromedially located gap. This calbindin-poor zone, termed the medial division of the inferior pulvinar (PIM), corresponds precisely to a region that contains elevated cytochrome oxidase activity and parvalbumin immunostaining. The PIM extends slightly above and behind the classically defined limit of the inferior pulvinar, the corticotectal tract. Regions of inferior pulvinar with intense immunostaining for calbindin-D28k were the posterior division of the inferior pulvinar (PIP, medial to PIM) and the central division (PIC, lateral to PIM). A newly recognized lateral region, PIL, adjoins the lateral geniculate nucleus and stains more lightly for calbindin and parvalbumin immunoreactivity and for cytochrome oxidase. Staining patterns for calbindin, parvalbumin, and cytochrome oxidase in the pulvinar of rhesus monkeys closely resemble those shown in squirrel monkey inferior pulvinar, suggesting that a common organization exists in all primates. In order to examine cortical connection patterns of the histochemically defined compartments in the inferior pulvinar, injections of up to five neuroanatomical tracers (wheat germ agglutinin conjugated to horseradish peroxidase and fluorescent retrograde tracers) were placed in the same cerebral hemisphere. Single injection sites were in the middle temporal area (MT), and several separate injections were placed in a strip corresponding to the rostral subdivision of the dorsolateral area (DLr). Injections that involved only DLr and not MT labeled principally the PIC, and more sparsely PIP and PIL. DLr connections occupied a "shell" region dorsal to PIM that extended from PIC into the lateral and medial divisions of the pulvinar, PL and PM. Injection sites that included MT or were largely restricted to MT produced dense label in PIM and moderate label in PIC and PIL. The retinotopic organization within the inferior pulvinar was inferred from patterns of connections. Connections with cortex related most closely to central vision were found posteriorly in PIM and in adjacent portions of PIC as it wraps around the caudal pole of PIM. Cortex related to more peripheral locations in the lower visual field connected with more rostral PIM and PIC. Patterns of label within the portions of PL and PM that were immediately adjacent to PIM roughly paralleled those in PIM and PIC.(ABSTRACT TRUNCATED AT 400 WORDS)

Protein kinase C-delta in rat brain: association with sensory neuronal hierarchies.

Originally characterized as the calcium- and phospholipid-dependent protein kinases, the protein kinases C include at least eight separate isoforms, some of which are calcium-independent and all of which are highly enriched in brain. Of the calcium-independent isoforms, the delta subspecies of protein kinase C has the most restricted complement of lipid activators and substrate specificity, suggesting that it may have a unique role in cell signalling pathways. Using immunocytochemistry, we report that the distribution of protein kinase C-delta immunoreactivity in rat brain is also restricted, being present in all sensory systems. Moreover, it is found in alternating hierarchies of sensory pathways: in all sensory systems except auditory, it is found in first- and third-order neurons, while in the auditory system, it is found in second- and fourth-order neurons. Thalamocortical systems are intensely immunoreactive, including barrel fields of the rat parietal cortex. Outside of sensory systems, protein kinase C-delta is present in cerebellum within longitudinal stripes in Purkinje neurons, and in the caudate-putamen, it appears to be associated with the striosome (patch) compartment. In contrast to all other protein kinase C isoforms, protein kinase C-delta is absent from hippocampus. These findings suggest that protein kinase C-delta may have a unique role in signal transduction in the central nervous system (CNS), especially in sensory systems.

Chronic deafferentation in monkeys differentially affects nociceptive and nonnociceptive pathways distinguished by specific calcium-binding proteins and down-regulates gamma-aminobutyric acid type A receptors at thalamic levels.

Chronic deafferentation of skin and peripheral tissues is associated with plasticity of representational maps in cerebral cortex and with perturbations of sensory experience that include severe "central" pain. This study shows that in normal monkeys the nonnociceptive, lemniscal component of the somatosensory pathways at spinal, brainstem, and thalamic levels is distinguished by cells and fibers immunoreactive for the calcium-binding protein parvalbumin, whereas cells of the nociceptive component at these levels are distinguished by immunoreactivity for 28-kDa calbindin. Long-term dorsal rhizotomies in monkeys lead to transneuronal degeneration of parvalbumin cells at brainstem and thalamic sites accompanied in the thalamus by a down-regulation of gamma-aminobutyric acid type A receptors and an apparent increase in activity of calbindin cells preferentially innervated by central pain pathways. Release from inhibition and imbalance in patterns of somatosensory inputs from thalamus to cerebral cortex may constitute subcortical mechanisms for inducing changes in representational maps and perturbations of sensory perception, including central pain.

Nerve injury-induced depletion of tachykinin immunoreactivity in the somatosensory cortex of adult squirrel monkeys.

Cortical immunoreactivity for tachykinin neuromodulators was evaluated 5-21 days after median and ulnar nerve transections in adult squirrel monkeys. Contralateral to the deafferentations, immunoreactive cells were reduced by about 20-40% in layer IV of the hand representation compared to the face region in area 3b. Similar deafferentations have been shown to alter the pattern of neuronal activity in the somatosensory cortex. Unresponsive regions are produced that with time become reactivated by receptive fields served by intact nerves. The immunocytochemical results reported here suggest that the presence or pattern of somatosensory input regulates content of tachykinin neuropeptides in intrinsic cortical neurons.

A sympathoinhibitory area in cat rostral medulla: its role in cardiovascular tone regulation and baroreflex.

We have found a medullary vasodepressor area in cat centered 3 mm rostral to the obex and just lateral to the compact division of the ambiguus nucleus. The area is compact, extending at most 1 mm in any direction. Microinjection of L-glutamate into this rostral depressor area (RDA) elicited acute hypotension and bradycardia. These responses were not reduced by either peripheral atropine blockade or bilateral vagotomy, but they were nearly abolished by peripheral phentolamine/propranolol blockade or high cervical cord transection. Bilateral reversible blockade of the RDA by local microinjection of the neuronal hyperpolarizing agent muscimol yielded chronic hypertension and tachycardia. Sympathetically mediated baroreflex, observed as a bradycardic response to a peripherally administered phenylephrine bolus in atropinized animals, was partially (50%) abolished during this same blockade. We conclude that the RDA contains sympathoinhibitory cells which are involved in the regulation of cardiovascular tone and in the expression of the sympathetic component of baroreflex.

Cortical connections of the caudal subdivision of the dorsolateral area (V4) in monkeys.

Evidence suggests that all primates have rostral and caudal subdivisions in the region of visual cortex identified as the dorsolateral area (DL) or V4. However, the connections of DL/V4 have not been examined in terms of these subdivisions. To determine the cortical connections of the caudal subdivision of DL (DLC) in squirrel monkeys, injections of the neuroanatomical tracers wheat germ agglutinin conjugated to horseradish peroxidase, Diamidino Yellow, and Fluoro-Gold were made in cortex rostral to V II. To aid in delineating the borders of DLC, cortex was also evaluated architectonically. Based on similar patterns of connections, DLC extends from dorsolateral to ventrolateral cortex. DLC receives strong, feedforward input from V II and projects in a feedforward fashion to the rostral subdivision of DL (DLR) and caudal inferior temporal (IT) cortex, including a separate location in the inferior temporal sulcus. DLC has weaker connections with V I, the middle temporal area (MT), cortex rostral to MT in the location of the fundal superior temporal area (FST), cortex dorsal to DLC, ventral cortex rostral to V II, and cortex in the frontal lobe, lateral to the inferior arcuate sulcus. Only lateral DLC has connections with V I, and only dorsolateral DLC has connections with cortex dorsal to DLC. The topographic organization of DLC was inferred from its connections with V II. Thus, dorsolateral DLC represents the lower field, lateral DLC represents central vision, and ventrolateral DLC represents the upper field. Limited observations were made on DLR. Confirming earlier observations (Cusick and Kaas: Visual Neurosci. 1:211, 1988), DLR is paler than DLC myeloarchitectonically. DLR receives only sparse feedforward input from V II, but stronger input from DLC. DLR has strong connections with cortex just rostral to dorsal V II, ventral posterior parietal cortex in the sylvian fissure, MT, the medial superior temporal area, FST, and the inferior temporal sulcus. DLR also shares connections with IT cortex. Thus, while both DLC and DLR are involved in the pathway relaying visual information to IT cortex, an area specialized for object vision, DLR also projects densely to areas such as MT involved in the pathway relaying to posterior parietal cortex, a region specialized for spatial localization and motion perception.

Cortical connections of dorsal cortex rostral to V II in squirrel monkeys.

A region of dorsal cortex along the rostral border of V II has been described as comprising a visual area or areas separate from more lateral cortex in both New and Old World primates. To evaluate these possibilities in squirrel monkeys, we studied patterns of cortical connections by injecting Fast Blue, Fluoro-Gold, horseradish peroxidase, and wheat germ agglutinin conjugated to horseradish peroxidase into the dorsal region and related results to distinctions in myeloarchitecture. Our major conclusions are as follows. 1) The dorsal region (D) has distinctly different connections from the area found laterally, the caudal subdivision of the dorsolateral area (DLC). These include major connections with the rostral subdivision of the dorsolateral area (DLR), ventral posterior parietal cortex in the Sylvian fissure, the middle temporal area (MT), the medial superior temporal area (MST), ventral cortex just rostral to V II, and cortex in the inferior temporal sulcus. Weaker connections are with V I, V II, DLC, the fundal superior temporal area (FST), and the frontal lobe. In contrast, DLC has strong connections with V II and inferior temporal (IT) cortex, weaker connections with DLR, and lacks connections with ventral posterior parietal cortex (Steele et al: J Comp Neurol 306:495-520, 1991). 2) Caudal and rostral aspects of dorsal cortex differ in the magnitude of connections with V I, V II, DLR, and FST. These differences are consistent with the previous proposal that at least two visual areas, caudal and rostral, occupy the dorsal region in squirrel monkeys (Krubitzer and Kaas: Visual Neurosci 5:165, 1990), but they could also reflect regional differences in the connections of a single visual area. 3) The dorsal region is more densely myelinated than surrounding cortex; however, rostral aspects of dorsal cortex are less myelinated than caudal aspects, again suggesting the existence of at least two areas. 4) The distinctiveness of connections between dorsal cortex and rostral as compared to caudal dorsolateral cortex provides further evidence for dividing the region of DL into two visual areas, DLC and DLR (Cusick and Kaas: Visual Neurosci 1:211, 1988; Steele et al: J Comp Neurol 306:495-520, 1991).

Temporal progression of cortical reorganization following nerve injury.

Damage to peripheral nerves of adult mammals causes reorganization of somatosensory maps in the cerebral cortex. An understanding of the temporal progression of cortical changes is important for understanding the underlying mechanisms. The present experiments utilized neurophysiological recordings to analyze the time course of reorganization in the S-I cortical hindpaw area in adult rats. Following loss of sciatic inputs, the cortical area responding to low threshold inputs from the hindpaw saphenous nerve expands. A brief, early onset period of rapid expansion is followed by a prolonged period of slow increase. The temporal progression suggests that early onset changes condition the central nervous system for later changes.

Connections between area 3b of the somatosensory cortex and subdivisions of the ventroposterior nuclear complex and the anterior pulvinar nucleus in squirrel monkeys.

The goal of this study was to determine whether somatosensory thalamic nuclei other than the ventroposterior nucleus proper (VP) have connections with area 3b of the postcentral cortex in squirrel monkeys. Small injections of the anatomical tracers wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP) or 3H-proline were placed in electrophysiologically identified representations of body parts. The results indicate that, besides the well-established somatotopically organized connections with VP, area 3b has connections with three other nuclei of the somatosensory thalamus: the ventroposterior superior nucleus (VPS ["shell" of VP]), the ventroposterior inferior nucleus (VPI), and the anterior pulvinar nucleus (Pa). Injections confined to area 3b or involving adjacent parts of area 3a or area 1 indicate that connections between VPS, VPI, and Pa and the postcentral cortex are somatotopically organized. In VPS, connections related to the hand were found medially, and connections related to the foot were lateral. In VPI, connections with the cortical representations of the mouth, hand, and foot were successively more lateral. In Pa, connections related to the mouth, hand, and foot were successively more ventral, lateral, and caudal, and the trunk region was caudomedial. The findings suggest that VPI contains a representation of all parts of the body, including the face. The connections of Pa with the primary somatosensory cortex, area 3b, the location of Pa relative to the ventroposterior nucleus, and the high degree of topographic order in the connections of Pa with the postcentral cortex suggest that Pa is an integral part of the somatosensory thalamus in monkeys and is homologous to the medial nucleus of the posterior group (Pom) in other mammals. Overall, the results contribute to the growing evidence that individual somatosensory cortical areas in monkeys receive inputs from multiple thalamic sources, and that a single thalamic nucleus has several cortical targets.