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H D Schwark

Publications and source records attributed to H D Schwark.

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Distribution and proportions of GABA-immunoreactive neurons in cat primary somatosensory cortex.

Certain receptive field properties of cortical neurons depend upon inhibitory, GABAergic inputs. In the somatosensory cortex, iontophoresis of bicuculline, a GABAA receptor blocker, results in enlargement of receptive fields. However, bicuculline's effectiveness in changing receptive field size varies with the neuron's adaptation characteristics, location within a particular submodality region, and laminar location. To test whether regional differences in the effectiveness of bicuculline are correlated with the distribution of cortical GABAergic neurons, we determined the numbers and proportions of GABA-immunoreactive [GABA(+)] neurons within cat primary somatosensory cortex. The laminar distribution of GABA(+) neurons was similar across all four cytoarchitectonic areas of primary somatosensory cortex, with layer II containing the highest areal density of GABA(+) neurons. Numerical proportions of GABA(+) neurons in the total neuron population were similar in areas 3b and 2 (29.8% and 22.6%, respectively). Laminar distributions of the proportions of GABA(+) neurons were also similar in these two areas; in both areas, layer I contained the highest proportion of GABA(+) neurons. The laminar distributions of GABA(+) neuron densities as well as GABA(+) neuron proportions differed from the reported laminar distribution of bicuculline effects on receptive field size. Moreover, within area 3b, these measures showed no evident patterns that might correspond to rapidly adapting and slowly adapting submodality regions.

Animals

Regional distribution of GABAA receptor binding sites in cat somatosensory and motor cortex.

Inhibition in primary sensory cortex plays a role in neuronal responses to peripheral stimuli. For many neurons in cat primary somatosensory cortex, blockade of GABAA receptors by bicuculline results in receptive field enlargement. The magnitude of this effect varies with the neuron's adaptation characteristics and its location in particular laminae and submodality regions. To test whether these variations are correlated with the distribution of GABAA receptors, we analyzed [3H]muscimol binding in cat primary somatosensory and motor cortical areas. The highest levels of binding were in layers I-III, and the lowest levels were in layers V-VI. In somatosensory cortical areas, layer IV was distinguished by higher levels of binding than in adjacent layers. Within layer IV, levels of binding were significantly higher in posterior area 3b than in anterior area 3b. These differences may correspond to the rapidly adapting and slowly adapting submodality regions which have been described in this area. The laminar distribution of [3H]muscimol binding differed from that of [3H]flunitrazepam, and neither resembled the distribution of the magnitude of bicuculline's effects on receptive field size. The laminar distribution of [3H]muscimol binding was highly correlated with the areal density of GABA-immunoreactive neurons described in a companion study.

Animals

Responses of rapidly adapting neurons in cat primary somatosensory cortex to constant-velocity mechanical stimulation.

1. The responses of rapidly adapting (RA) neurons to constant-velocity ramp stimulation were studied in the forepaw representation of primary somatosensory cortex (SI) of anesthetized cats. Single probe stimuli were used to indent the skin or to move hair parallel to the skin surface. The velocity of the moving stimulus probe was varied to determine the rate sensitivity of the neurons. 2. The cortical RA neurons were classified into four categories identified as G1/F1, Gint/Fint, G2/F2, and complex classes. The primary bases for classification in the present experiments were the pattern of response during ramp stimulation, velocity threshold, and directional sensitivity. 3. Of the RA neurons recorded in SI, 84% (49/58) could be assigned to one of the three response classes with little ambiguity. The remaining neurons showed more complex responses. The form of the complex responses suggested that they arose from a combination of inputs of different response classes. Some of these appeared to arise from a combination of different RA input classes, whereas others had components that resembled responses previously described for C mechanoreceptors. 4. Increased ramp velocity resulted in increased average firing frequency in 87% of the RA neurons. This relationship, which could be fitted with a power function, varied with response class. G1/F1 neurons were more sensitive to stimulus rate than G2/F2 neurons. Significant differences between response classes also were seen in the relationship between ramp velocity and their number of evoked action potentials and in their spontaneous firing rates. 5. The results demonstrate that a discrete SI neuron population is sensitive to the rate of stimulus movement. This observation is consistent with psychophysical studies reporting effects of stimulus indentation rates on perception of single probe stimuli. The appearance of complex responses in a small proportion of SI neurons provides evidence of convergence in somatosensory pathways to SI.

Afferent Pathways

Receptive field reorganization in dorsal column nuclei during temporary denervation.

Altered sensory input can result in the reorganization of somatosensory maps in the cerebral cortex and thalamus, but the extent to which reorganization occurs at lower levels of the somatosensory system is unknown. In cat dorsal column nuclei (DCN), the injection of local anesthetic into the receptive fields of DCN neurons resulted in the emergence of a new receptive field in all 13 neurons studied. New receptive fields emerged rapidly (within minutes), sometimes accompanied by changes in adaptation rates and stimulus selectivity, suggesting that the new fields arose from the unmasking of previously ineffective inputs. Receptive field reorganization was not imposed by descending cortical inputs to the DCN, because comparable results were obtained in 10 additional cells when the somatosensory and motor cortex were removed before recording. These results suggest that mechanisms underlying somatotopic reorganization exist at the earliest stages of somatosensory processing. Such mechanisms may participate in adaptive responses of the nervous system to injury or continuously changing sensory stimulation.

Animals

Distribution of [3H]QNB and [125I]alpha-bungarotoxin binding and acetylcholinesterase activity in visual system and hippocampal structures of eleven mammalian species.

This study assessed interspecies differences in regional brain distribution of [3H]QNB binding, [125I]alpha-bungarotoxin binding and acetylcholinesterase activity, by autoradiographic and histochemical methods. Eleven mammalian species were examined, including carnivores (cat, dog), a lagomorph (rabbit), and rodents (squirrel, guinea pig, gerbil, hamster, vole, lemming, rat, mouse). Comparisons were based on primary visual system structures (superior colliculus, lateral geniculate nucleus, primary visual cortex) and the hippocampal formation. The two radioligands differed greatly in the degree of interspecies variation: while the pattern of [3H]QNB binding was quite similar across species, [125I]alpha-bungarotoxin showed striking interspecies diversity. This contrast was most obvious in laminar patterns of the visual cortex and hippocampal formation. Regional distributions of acetylcholinesterase staining were fairly diverse, and were unlike the patterns of either [3H]QNB or [125I]alpha-bungarotoxin. The two ligands showed more consistency in overall levels across species than did acetylcholinesterase. Possible correlates of the differences in interspecies diversity are discussed.

Acetylcholinesterase

Corticocortical connections of cat primary somatosensory cortex.

The organization of corticocortical connections in the representation of the forepaw in cat primary somatosensory cortex (SI) was studied following injections of various tracers into different cortical cytoarchitectonic areas. Small injections of horseradish peroxidase, wheat germ agglutinin-conjugated HRP, Phaseolus vulgaris leukoagglutinin, or fast blue were placed into the representation of the forepaw in areas 3b, 1, or 2. The positions of labeled neurons in SI and the surrounding cortical areas were plotted on flattened surface reconstructions to determine the organization of the corticocortical connections within SI. A strong, reciprocal projection linked the two forepaw representations which have been described in area 3b and the part of area 2 which lies in the anterior bank of the lateral ansate sulcus (see Iwamura and Tanaka 1978a, b). Dense projections also linked these areas with SII, as previously reported (Burton and Kopf 1984a). Additional projections to area 3b arose primarily from areas 3a and 1. Projections to area 2 were more widespread than those to area 3b, and arose from all other areas of SI as well as from areas 4 and 5a. All injections into SI tended to label groups of neurons which lay in mediolateral strips. Corticocortical projection neurons which were most heavily labeled by SI injections were pyramidal cells in layer III. Additional projections from area 2 to 3b, area 5a to 2, and SII to areas 2 and 3b arose from layer VI as well. Although neurons of layers III and VI were always the most densely labeled, large injections into SI labeled neurons in layers II and V as well.

Animals

The distribution of intrinsic cortical axons in area 3b of cat primary somatosensory cortex.

The morphology of single neurons in area 3b of cat primary somatosensory (SI) cortex was examined after horseradish peroxidase (HRP) injections. Neurons were labeled either by intracellular injection of HRP following intracellular recording or by small extracellular iontophoretic HRP injections. Both pyramidal and nonpyramidal neurons were labeled and reconstructed from serial sections. Their axons had local, interlaminar and interareal patterns of termination. Most neurons formed local axonal fields around their cell bodies and dendrites. Pyramidal neurons in cortical layer IV sent axons up into layers II and III, neurons in layers II and III sent axons down to layer V, and layer V neurons sent axons to layer VI as well as back to the upper layers. Layer VI neurons sent axons back to the upper cortical layers in a unique bowl-shaped pattern. The horizontal distribution of axons of pyramidal cells in layer III was extremely widespread. Axons of layer III neurons in area 3b terminated within 3b and area 1, but not in other areas of SI. Layer III neurons in area 1 distributed axon collaterals to all fields of SI as well as projecting a main axon to motor cortex. In general, the axon collaterals of area 3b pyramidal cells outside layer III remained confined to area 3b. Most of the nonpyramidal neurons labeled were basket cells in layers III and VI. These neurons formed dense axonal fields around their cell bodies, and none of their axons could be followed into the underlying white matter. The results of the present study demonstrate that area 3b somatosensory cortical neurons and their axons are vertically organized in a manner similar to that reported for other sensory cortical areas. They also show that widespread horizontal connections are formed by pyramidal neurons of layer III, and that these horizontal axons can travel for great distances in the cortical grey matter.

Animals

Morphology of physiologically characterized medial lemniscal axons terminating in cat ventral posterior thalamic nucleus.

1. Medial lemniscal axons were identified by extra- and intracellular recording in the thalamic ventral posterior lateral nucleus (VPL) of cats and injected intracellularly with horseradish peroxidase (HRP). 2. Axons were characterized in terms of their latencies of response to stimulation of the medial lemniscus in the medulla, their receptive fields, and the temporal patterns of their discharge in response to stimulation of the receptive field with natural, hand-held stimuli. One-hundred sixty-six axons were placed in five operational groups: hair transient (Ht) (n = 41); hair sustained (Hs) (n = 45); pressure transient (Pt) (n = 14); pressure sustained (Ps) (n = 27), and deep or joint (Jt) (n = 39). 3. There was a tendency for Jt axons to have their terminations in anterodorsal parts of VPL and for those in the four cutaneous categories to have theirs in more central parts of the nucleus. 4. Nineteen injected axons with receptive fields mainly on the distal forelimb were subjected to detailed morphological analysis in terms of extent of terminal field and number of boutons. All axons ended in localized terminal fields that were more extensive anteroposteriorly than in the other dimensions. All showed an overall similarity and similar ranges of variation. There was a tendency, however, for Jt axons to have the least extensive terminations with fewest boutons. Ps axons had the most extensive terminations and largest number of boutons; Hs axons had small terminations and few boutons but Ht axons had small-to-medium arborizations with many boutons; no Pt axons were sufficiently well stained to enable comparisons of them with the others. There were no marked differences in axon diameter or conduction velocity among the five types. 5. Boutons identified light microscopically tended to be clustered in linear chains along proximal dendrites of relay neurons and electron microscopy revealed that they were terminals making synaptic contacts on relay cell dendrites and on presynaptic dendrites of interneurons. 6. These results reveal more similarities than differences among lemniscal axon terminations in VPL. Further studies of a quantitative nature on stimulus-response coupling and on the geographic distribution of lemniscal synapses on relay neurons will be required to reveal how lemniscal input is translated into relay cell output in VPL.

Animals

Numbers and proportions of GABA-immunoreactive neurons in different areas of monkey cerebral cortex.

The number and proportion of neurons displaying GABA immunoreactivity were determined for 50-micron-wide columns through the thickness of 10 areas of monkey cerebral cortex, including the precentral motor area (area 4), 3 cytoarchitectonic fields of the first somatic sensory area (areas 3b, 1, and 2), 2 areas of parietal association cortex (areas 5 and 7), the first and second visual areas (areas 17 and 18), area 21 of the temporal lobe, and areas of the orbital and lateral frontal cortex. Methods of fixation and immunocytochemical processing were designed to maximize the number of stained cells in 15-micron-thick frozen sections and 1-micron-thick plastic sections. In 8 of the 10 areas the number and proportion of GABA-immunoreactive neurons per 50-micron-wide column were found to be the same (34-43 cells/column; 25% of the total neuronal population). Areas 17 and 3b differed. Area 17 contained 50% more GABA-immunoreactive neurons (52-66 cells/column) but more than twice the total number of neurons, so that the GABA cells made up less than 20% of the total. In 3 monkeys, the number and proportion of GABA-positive neurons per 50-micron-wide column in area 3b were smaller than in adjacent areas of sensorimotor cortex (26-42 cells/column; 19-22%). In 2 other monkeys, the number and proportion (34-43 cells/column; 24-26%) were the same as in adjacent areas. Despite the similarity among most areas of monkey cortex, within some areas, the number of GABA-positive neurons per 50-micron-wide column varied as much as 30%. These variations form a significant, repeating pattern only in area 18, where narrow bands (150-200 micron wide) of relatively few stained cells alternated with either narrow or wide bands (600-700 micron wide) in which columns contained more cells. The GABA-immunoreactive neurons were unevenly distributed across layers, with every area containing large numbers and proportions of stained cells in layer II, and every area but area 4 displaying a second concentration in the principal thalamocortical recipient layers. In area 4, the number of GABA-positive neurons declined sharply from layer II to layer III and remained low through layer VI. For areas displaying the greatest intra-areal variability, only 1 or 2 layers contributed significantly to that variability (layer IV in area 3b, layers III and V in area 18, and layers II and III in area 17).(ABSTRACT TRUNCATED AT 400 WORDS)

Animals

Extent of the ipsilateral representation in the ventral posterior medial nucleus of the monkey thalamus.

Single and multiunit mapping was used to determine the extent of the representation of ipsilateral structures in the ventral posterior medial (VPM) nucleus of the thalamus in cynomolgus monkeys. The extent of the VPM occupied by terminations of afferent fibers arising in the ipsilateral principal trigeminal nucleus was also determined by anterograde transport of horseradish peroxidase. Both methods indicate that most of the medial half of VPM is occupied by the ipsilateral representation. This is much larger than previously suspected. Units in the medial half of VPM have small, well localized receptive fields on the ipsilateral side of the lower lip, tongue and palate, in the ipsilateral cheek pouch and on the ipsilateral teeth. The representation is largest for the ipsilateral side of the tongue and the cheek pouch. Most units in the lateral half of VPM have small, contralateral receptive fields. Few units in VPM have bilateral receptive fields. VPM is clearly distinguishable by cytochrome oxidase (CO) staining. Anteroposteriorly elongated, CO-positive aggregations correspond to elongated aggregations of units with the same or closely similar receptive fields, especially in the medial, ipsilateral representation.

Animals

Cat area 17. I. Pattern of thalamic control of cortical layers.

Reversible inactivation of individual layers of the cat lateral geniculate and medial interlaminar nuclei was used to investigate the necessary and sufficient inputs for maintaining visually driven activity and receptive field properties in area 17. Neither orientation selectivity nor direction selectivity depends on any individual geniculate layer. We identified two groups of cortical layers on the basis of the pattern of thalamic inputs providing visual driving through the contralateral eye. One group, consisting of layers 4 and 6, has geniculate layer A as its only necessary and sufficient input. The other, consisting of supragranular layers, integrates at least two sufficient thalamic inputs, one of which is layer A. Several major receptive field properties are independently generated in these two groups of layers.

Animals

Cat area 17. II. Response properties of infragranular layer neurons in the absence of supragranular layer activity.

Response properties of cells in the infragranular layers of cortical area 17 of the cat were examined in the absence of input from supragranular layers. Supragranular activity was silenced either reversibly by cooling the surface of cortex or permanently by making a cryogenic lesion of the supragranular layers. Visually driven responses of cells throughout the cortical column were recorded with a linear array of electrodes. Most infragranular layer cells continued to be visually responsive in the absence of supragranular layer input. These cells were similar to normal infragranular layer cells on measures of visual responsiveness, orientation selectivity, and direction selectivity. Special complex, but not standard complex, cells were absent in layer 5 when supragranular layers were destroyed. We found no evidence for a selective effect of removal of supragranular activity on the response properties of cells in layer 6. We propose that the intracolumnar projection from the supragranular layers drives the special complex cells of layer 5, but is not necessary for the visual driving of most other infragranular layer cells. This projection does not impose selectivity for stimulus orientation or direction on the remaining active cells of the infragranular layers.

Animals

Cat area 17. III. Response properties and orientation anisotropies of corticotectal cells.

The receptive field properties of antidromically identified corticotectal (CT) cells in area 17 were explored in the paralyzed, anesthetized cat. To compare these with another population of infragranular cells, we also examined the receptive field properties of cells in layer 6. Sixty percent of our sample of CT cells showed increased response to increased stimulus length (length summation) and were classified as standard complex cells. The other 40% showed little or no length summation, were generally end stopped, and were classified as special complex cells. Standard and special complex CT cells have complementary orientation anisotropies: the distribution of orientation preferences of standard complex cells is biased toward obliquely oriented stimuli, whereas special complex cells are biased toward horizontally and vertically oriented stimuli. The receptive fields of the cells in our sample were primarily along the horizontal meridian so we cannot determine if these anisotropies are defined relative to the vertical meridian or relative to the meridian passing through the receptive field. The effects of these anisotropies in preferred orientation are minimized by the broad orientation tuning of CT cells. There was no simple relationship between the direction bias of CT cells and the reported direction bias of tectal cells. In contrast to the heterogeneity of corticotectal cells, layer 6 cells uniformly showed strong length summation, tight orientation tuning, and little spontaneous activity.

Animals

Cat area 17. IV. Two types of corticotectal cells defined by controlling geniculate inputs.

The dependence of cat area 17 corticotectal (CT) cells on specific subdivisions of the dorsal lateral geniculate (LGN) and medial interlaminar nuclei (MIN) was examined using reversible inactivation techniques. Inactivation of layer C of the LGN or layer 1 of the MIN did not block visual activity of CT cells driven through the contralateral eye. Inactivation of layer A of the LGN revealed two populations of CT cells: one strongly dependent on layer A and one whose visually driven activity survived layer A inactivation. CT cells that responded best to short stimuli (special complex cells) were least dependent on layer A, whereas cells that responded best to long stimuli (standard complex cells) were most dependent on layer A. We propose a model of the intracortical circuitry of these two types of CT cells. Standard complex cells, which are heavily dependent on layer A, receive sustaining visual input through layers 4 and/or 6. Special complex cells, which are not dependent on any single layer of the lateral geniculate nucleus, receive sustaining visual input from supragranular layers.

Animals

Cat medial interlaminar nucleus: retinotopy, relation to tapetum and implications for scotopic vision.

The medial interlaminar nucleus (MIN) of the cat was electrophysiologically mapped in sufficient detail to resolve individual laminae and to allow reconstruction of isoazimuth and isoelevation lines in coronal, sagittal, and horizontal planes. The electrophysiologically defined laminar pattern was in agreement with that revealed anatomically in the same animal, as well as with the general pattern revealed by anatomical methods in several unmapped nuclei. The MIN is made up of three distinct layers, each receiving inputs from one hemi-retina but none representing an entire hemifield. We confirm the findings of Guillery et al. (19) that the contralateral hemifield is represented in layers 1 and 2 through the contralateral and ipsilateral eyes, respectively, and that layer 3 represents the ipsilateral hemifield through the contralateral eye. Elevation and absolute value of azimuth are represented continuously through the MIN. When an isoazimuth line crosses the border between layer 3 and either layer 1 or 2, the absolute value of azimuth is maintained but the sign of the azimuth changes. Adjacent points on either side of this border represent mirror symmetrical visual directions on opposite sides of the vertical meridian. This indicates that the distance from the vertical meridian is an independently coded parameter within the geniculate complex. There is virtually no nasotemporal overlap in any layer of the MIN. The function relating magnification (mm3 per steradian) to eccentricity is strikingly similar to the function relating retinal ganglion cell density to eccentricity, suggesting that a constant fraction of retinal ganglion cells project to the MIN at all eccentricities. Most of the volume of each MIN layer is devoted to lower visual fields. Analysis of the geniculate retinotopic maps of Sanderson (46) reveals no equivalent bias toward lower visual fields in the dorsal lateral geniculate nucleus. The MIN represents a region of retina roughly coincident with the tapetum, suggesting a role of the MIN in dim-light vision.

Animals

Olfactory granule cell development in normal and hyperthyroid rats.

Dendritic development was examined in olfactory bulbs of both normal 7-, 14-, 21- and 60-day-old rats and littermates treated on postnatal days 1-4 with 1 microgram/g body weight of L-thyroxine sodium. Tissue was processed via the Golgi-Cox technique and subjected to quantitative analyses of mitral and internal layer granule cell development. These populations of granule cells were selected because their pattern of late proliferation suggested potentially greater susceptibility to postnatal hormonal alterations. Although neonatal hyperthyroidism induces widespread acceleration of maturation, including precocious chemosensitivity, granule cell development was unaffected relative to littermate controls. Both normal and hyperthyroid groups exhibited an inverted U-shaped pattern of cellular development, with rapid dendritic dendritic growth and expansion occurring during the earliest ages tested, but with loss of processes and dendritic field size occurring after day 21.

Aging