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U T Eysel

Publications and source records attributed to U T Eysel.

At least 19 recordsLinked to original sources

Effect of sensory deafferentation on immunoreactivity of GABAergic cells and on GABA receptors in the adult cat visual cortex.

To investigate the effects of sensory deafferentation on the cortical GABAergic circuitry in adult cats, glutamic acid decarboxylase (GAD) and gamma-aminobutyric acid (GABA) immunoreactivity and GABA receptor binding were studied in the visual cortex of normal cats and compared with cats that had received restricted binocular central lesions of the retina and had survived for 2 weeks postlesion in a normal visual environment. In the visual cortex of lesioned cats, two changes were observed in the number of GAD-immunoreactive elements in the regions affected by the retinal lesions: the number of GAD-positive puncta decreased, whereas that of GAD-immunoreactive somata increased. In contrast, no detectable changes were measured in the number of GABA-immunopositive somata or puncta. At the receptor level, we observed no differences in either the laminar distribution or the affinity of cortical GABAA and GABAB receptors labeled with [3H]-muscimol and [3H]-baclofen, respectively, in the lesioned versus normal cats. We present the hypothesis that sensory deafferentation in these adult cats (1) leads to a reduction of cortical GABAergic inhibition in the deafferented region, and (2) that this decreased inhibition may permit changes in efficiency of synapses and (3) that these changes may represent a first stage of events underlying the retinotopic reorganization preceeding the structural changes.

Afferent Pathways

Possible enhancement of GABAergic inputs to cat dorsal lateral geniculate relay cells by serotonin.

The possible enhancement of inhibitory inputs to relay cells of the dorsal lateral geniculate nucleus (dLGN) by serotonin (5-HT) was studied during blockade of GABAA-receptor mediated inhibition with bicuculline (BICU). Visually induced spike activity of single units was recorded during simultaneous micro-iontophoretic application of BICU or 5-HT. All 28 relay cells studied were disinhibited by BICU and strongly inhibited by 5-HT. The responses during BICU or 5-HT were on average statistically different from the controls (p < 0.01, Student's t-test). The inhibition by 5-HT could be almost totally abolished by the simultaneous application of BICU and responses during the combined application of both substances were only slightly different from those obtained during BICU alone (p < 0.3).

Animals

Time-slice analysis of inhibition in cat striate cortical neurones.

A prominent feature of visual cortical neurones is orientation specificity. The underlying mechanisms and the possible contributions of intracortical inhibition are continuously the subject of intense debate. We analysed orientation tuning in relation to time after stimulus onset and the effects of blockade of GABAergic inhibition with microiontophoretically applied bicuculline in single cells in area 17. The majority of the cells had sharp and temporally stable orientation specificity; however, changes of receptive field structure, widening of orientation tuning and changes in preferred orientation were observed between 30 and 150 ms after stimulus onset when inhibition was blocked. This indicates that under normal conditions in most in most striate cortical cells intracortical inhibition sculptures receptive field structure and keeps orientation tuning narrow and constant in time.

Animals

Relationship between lateral inhibitory connections and the topography of the orientation map in cat visual cortex.

The functional and structural topography of lateral inhibitory connections was investigated in visual cortical area 18 using a combination of optical imaging and anatomical tracing techniques in the same tissue. Orientation maps were obtained by recording intrinsic signals in regions of 8.4-19 mm2. To reveal the inhibitory connections provided by large basket cells, biocytin was iontophoretically injected at identified orientation sites guided by the pattern of surface blood vessels. The axonal and dendritic fields of two retrogradely labelled large basket cells were reconstructed in layer III. Their axonal fields extended up to 1360 microns from the parent somata. In addition to single basket cells, the population of labelled basket cell axons was also studied. For this analysis anterogradely labelled basket axons running horizontally over 460-1280 microns from the core of an injection site in layer III were taken into account. The distribution of large basket cell terminals according to orientation preferences of their target regions was quantitatively assessed. Using the same spatial resolution as the orientation map, a frequency distribution of basket cell terminals dependent on orientation specificity could be derived. For individual basket cells, the results showed that, on average, 43% of the terminals provided input to sites showing similar orientation preferences (+/- 30 degrees) to those of the parent somata. About 35% of the terminals were directed to sites representing oblique-orientation [+/- (30-60) degrees], and 22% of them terminated at cross-orientation sites [+/- (60-90) degrees]. Furthermore, the possible impact of large basket cells on target cells at different distances and orientation preferences was estimated by comparing the occurrence of orientation preferences with the occurrence of basket terminals on the distance scale. It was found that a basket cell could elicit iso-orientation inhibition with a high impact between 100-400 and 800-1200 microns, strong cross-orientation inhibition at approximately 400-800 microns, and oblique-orientation inhibition between 300-500 and 700-900 microns from the parent soma. The non-isotropic topography of large basket axons suggests a complex function for this cell class, possibly including inhibition related to orientation and direction selectivity depending on the location of the target cells and possible target selectivity.

Animals

Transient immunohistochemical labelling of rat retinal axons during Wallerian degeneration by a monoclonal antibody to neurofilaments.

Immunohistochemical labelling with the monoclonal antibody SMI32 to non-phosphorylated epitopes on neurofilament proteins of high molecular weight class was low in rat central optic fibers of controls. After unilateral transection of optic nerve, a strong, transient increase of labelling with SMI32 occurred in degenerating fibers of optic tract at 2 and 4 days, which then declined at 8 and remained low at 21 days. Consequently, immunostaining with SMI32 may serve as a positive marker for degenerating fibers in rat optic system.

Animals

Alterations in GAP-43 and synapsin immunoreactivity provide evidence for synaptic reorganization in adult cat dorsal lateral geniculate nucleus following retinal lesions.

Growth-associated protein-43 (GAP-43) and synapsin were used as molecular markers for synaptic reorganization in the adult cat visual system following sensory deprivation. Small binocular retinal lesions (central 10 degrees) were made with a xenon light photocoagulator in adult cats. One, 3, 5 and 7 weeks after induction of the lesion, the neuropil levels of synapsin and GAP-43 in the dorsal lateral geniculate nucleus (dLGN) and area 17 were determined by immunocytochemistry. GAP-43 displayed a moderately low basal level in the dLGN of normal adult cats. The parvocellular C layers and the interlaminar plexi were characterized by higher immunoreactivity for GAP-43. Lesion-induced alterations were observed in all layers: GAP-43 immunoreactivity increased in the part of the dLGN representing central vision. This increase was maximal 3 weeks after the lesion. Under our experimental conditions, sensory deprivation did not significantly alter GAP-43 levels in the visual cortex. The changes in synapsin immunoreactivity were also restricted to the dLGN. In this nucleus, synapsin immunoreactivity decreased in all layers in the part subserving central vision 1 week after lesion. By 3 weeks after lesion, the level of synapsin had already returned to normal. This study provides evidence for a capacity for structural remodelling in primary sensory brain areas such as the dLGN throughout adult life. The observed changes in GAP-43 and synapsin in the dLGN suggest that synaptic reorganization is induced by retinal lesions. Normalization of synaptic density and activity could be important for the survival of the partially deafferented geniculate neurons.

Animals

Velocity invariance of preferred axis of motion for single spot stimuli in simple cells of cat striate cortex.

Directional tuning for motion of a long bar and a spot was compared quantitatively over a wide range of velocities in 23 simple cells of cat striate cortex whose "on" and "off" receptive field subregions had been mapped with optimally oriented, stationary flash-presented bars. Tuning curves were derived using stimuli whose polarity of contrast was appropriate for the dominant receptive field subregion of each cell (i.e. light stimuli for on-subregions and dark stimuli for off-subregions); stimulus sweep was centred accurately on the centre of that subregion. Bar stimuli were of optimal width, and spot diameter was equal to the width of the bars. In all simple cells, preferred axis of motion for a long bar was invariant with velocity, being orthogonal to preferred orientation, as assessed with a stationary flash-presented bar. In 20 of 23 simple cells, preferred axis for spot motion was approximately orthogonal to that for bar motion (i.e., parallel to preferred orientation) at all velocities tested, including those just above threshold for spot stimuli. However, tuning for the spot became sharper as velocity was increased, due to an increase in response to the spot moving along the preferred axis and a decrease in response to spot motion along other axes, including the preferred axis for the bar. Both preferred and upper cut-off velocity were consistently higher for spot than for bar motion. The remaining 3 simple cells showed no response to spot motion at any velocity, and their preferred axis of motion for the shortest bar which evoked a consistent response was the same as the long bar.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals

Lesion-induced transient suppression of inhibitory function in rat neocortex in vitro.

The structural and functional consequences of a local thermolesion were examined in rat neocortex with electrophysiological in vitro techniques and immunocytochemistry. Age-matched untreated and sham-operated animals served as controls and were analysed in the same way. The lesions consisted of a core of coagulated tissue 2-3 mm in diameter and reached ventrally into the deep cortical layers. After two days reactive astrocytes and after nine days a dense gliosis were observed in the immediate vicinity. Modifications in the intrinsic membrane characteristics and the synaptic network properties were investigated with intra- and extracellular recording techniques after survival times of one to eight days. Neurons recorded in the surrounding of lesions in neocortical slices revealed a significantly more depolarized resting membrane potential and a higher neuronal input resistance. In comparison to cells in control slices, maximal discharge rates to injection of depolarizing current pulses of neurons close to a focal lesion were not significantly altered and intrinsic burst firing was never observed. However, between postlesion days 1 and 5, neurons in the surroundings of lesions showed a transient increase in synaptic excitability. This hyperactivity was most clearly pronounced at a distance of 2-3 mm from the centre of the lesion (i.e. about 1-1.5 mm away from the lesion border) and characterized by long-duration field potential responses and multiphasic long-lasting excitatory postsynaptic potentials to orthodromic stimulation of the afferent input. This lesion-induced hyperexcitability was associated with a significant reduction in the peak conductance of the Cl(-)-dependent fast inhibitory postsynaptic potential and the K(+)-dependent long-latency inhibitory postsynaptic potential, suggesting that the intracortical GABAergic system was functionally impaired. The decrease in synaptic inhibition was associated with prolonged N-methyl-D-aspartate receptor-mediated activity, which could be reversibly blocked by D-amino-phosphonovaleric acid. In addition, neurons recorded in the vicinity of the lesion responded to an orthodromic synaptic stimulus with a long-lasting burst. The lesion-induced disturbance in the balance between the excitatory and inhibitory system may not only have profound influences on the mechanisms of intracortical information processing, but may also lead to the expression of epileptiform activity and long-term functional deficits.

Action Potentials

Ischemic damage visualized in flat mounts of rat retina after photochemically induced thrombosis.

The method of photochemically induced thrombosis was used to produce severe ischemia in the rat retina. Flat mounts of the retina were prepared at 3 h, and 1, 2, 3, 4, 6, 7, and 22 days after lesioning and Nissl-stained, which facilitated the study of the topography of the ischemic lesions. The regional variability of ischemic damage and the cytological features of ischemic cell death in the ganglion cell layer were evaluated. Neuropathological analysis showed ischemic cell damage of ganglion cells at 3 h, an infarction-type lesion at 1 to 7 days, and scar formation at 3 weeks. As an additional parameter of ischemic ganglion cell death, the degeneration of retinal axons was visualized in the contralateral dorsal lateral geniculate nucleus by Fink-Heimer silver impregnation and by immunohistochemical staining for glial fibrillary acidic protein (GFAP) in reactive astrocytes.

Animals

[Increased spontaneous and evoked response in the area of model infarcts of the visual cortex of cats].

Ischemic lesions were produced in the cat visual cortex and changes of responses to visual stimuli were studied in single cells with extracellular recordings. The spontaneous and the visually driven activity were quantitatively analyzed and compared with those in normal controls. While the normal control cells showed a mean response strength of 52.7 impulses/s, this value was increased to 69.3 impulses/s in the vicinity of ischemic lesions. The spontaneous activity was also increased. Epileptiform burst activity with frequencies above 500 I/s was observed in 37.7% of the cells. Directional and orientational specificity was significantly reduced in these regions because of an increase in responses to stimuli in non-preferred directions and with non-preferred orientations. Thus, an increase in activity-dependent energy consumption occurs in the vicinity of lesions in the ischemically severed cortex.

Animals

Functional and structural topography of horizontal inhibitory connections in cat visual cortex.

The functional organization of long-horizontal inhibitory connections was studied in cat visual cortical area 17, using a combination of electrophysiological recording and anatomical tracing in the same tissue. Orientation maps were obtained by recording multiunit activity from layer III at regular intervals (100-300 microns) in a region of approximately 1.3 mm2 of cortex at a depth corresponding to the location of the basket cell axons reconstructed later. Before the physiological mapping, the neuronal tracer biocytin had been iontophoretically injected at one functionally characterized site. On the basis of light microscopic features a total of five biocytin-labelled large basket axons, BC1-BC5, were reconstructed from series of horizontal sections of two cats. The parent somata and dendritic fields of three axons (BC1, BC4 and BC5) could also be reconstructed. The axonal field of basket cell BC1 had an overall lateral spread of 1.8 mm. The axons of basket cells BC4 and BC5 spanned a distance of 3.05 and 2.85 mm, respectively. The distribution pattern of histologically reconstructed recording sites and of five labelled basket cell axons were directly compared in the same sections. The results show that a single large basket cell provides input to regions representing the whole range of orientations, i.e. iso-orientation (+/- 30 degrees), oblique orientation (+/- [30-60] degrees) and cross-orientation (+/- [60-90] degrees) to that at the basket cell's soma. Furthermore, the differential effect mediated by the same large basket cell at sites of different orientation preference was numerically estimated for two basket cells (BC4 and BC5) whose preferred orientations could be determined on the basis of recording sites adjacent to their parent somata. We counted the number of axonal terminals of these basket cells at iso-, oblique- and cross-orientation sites and found no significant difference in the average density of terminals at sites of either orientation preference. The functional topography of large basket cell axons indicates that the same basket cell can mediate iso-, oblique- and cross-orientation inhibition at different sites. Hence, we assume that large basket cells serve a complex physiological role depending on the location of target cells in the orientation map.

Animals

Network of GABAergic large basket cells in cat visual cortex (area 18): implication for lateral disinhibition.

Anatomical and immunohistochemical data indicate that, in addition to pyramidal neurons, nonpyramidal cells are exposed to perisomatic inhibition mediated by gamma-aminobutyric acid (GABA)-containing terminals. However, no direct information is available as yet for the origin of GABAergic inputs to morphologically identified GABAergic neurons. In the present paper, we studied the topographical and synaptic relationship between identified GABAergic large basket cells and their immunohistochemically characterized target neurons revealed by parvalbumin-(PV) and GABA immunostaining in the same material. Extracellularly applied biocytin labelled a total of 36 and 9 large basket cells in layers III and V, respectively. Of these, the axonal arborizations of two basket cells, BC1 and BC2, were reconstructed. The axon of BC1 occupied an area of about 2.3 x 2.2 mm2 in layer III, providing a total of 2,755 terminals. The axon of BC2 showed an overall extent of 3.8 x 1.7 mm2 in layer V elongated in the anteroposterior direction, and gave off 1,599 terminals. Immunostaining for PV was carried out to reveal putative nonpyramidal targets for BC1 and BC2. It was found that in addition to immunonegative cells, they established an average of 4-6 perisomatic contacts onto each of 58 (BC1) and 33 (BC2) PV-immunopositive neurons. For electron microscopic verification, 23 terminals apposing the somata of 12 PV-immunopositive neurons were selected. Each terminal was found to establish symmetrical (type II) contacts with its targeted cell. Furthermore, the distribution of soma area of the targeted PV-immunopositive cells and of identified large basket cells showed remarkable similarity, implying that the two populations were actually the same. In addition, the average horizontal distance between neighbouring PV-immunopositive target cells was found to be about 100 microns both in layers III and V. The results suggest that in area 18 the same large basket cell provides direct inhibition to certain pyramidal cells and facilitation to other pyramidal neurons, by inhibiting their presynaptic large basket cells at regular intervals.

Animals

Modulatory effects of acetylcholine, serotonin and noradrenaline on the activity of cat perigeniculate neurons.

We studied the modulatory actions of microiontophoretically applied acetylcholine (ACH), serotonin (5-HT, 5-hydroxytryptamine) and noradrenaline (NA), and those of the adrenoceptor agonists phenylephrine (PHE, alpha 1), clonidine (CLO, alpha 2) and isoprenaline (ISO, beta) on spontaneous and visually induced activities in cat perigeniculate (PGN) and thalamic reticular (NRT) neurons (only spontaneous) during extracellular recordings performed in vivo. ACH and 5-HT were found to affect the ongoing (spontaneous) and visually evoked activity of PGN cells and also the spontaneous activity of NRT cells in an opposite fashion. ACH inhibited tonic firing and often induced burst activity. By contrast, 5-HT exerted an excitatory influence, which caused a long-lasting, very regular, high-frequency activity between about 35 and 120 Hz. Spontaneous as well as 5-HT-induced firing was found to prefer three distinct frequency ranges: 35-42 Hz, 60-67 Hz and 80-120 Hz. Opposite actions of ACH and 5-HT were also evident when applied simultaneously. ACH dampened the high-frequency activity elicited with 5-HT, and 5-HT could replace the burst activity induced with ACH application by a regular tonic activity. The absolute strength of visual responses (in spikes per second) was only slightly enhanced or reduced by ACH and 5-HT, respectively, but due to the strong effects on background activity, ACH clearly elevated the signal-to-noise ratio and 5-HT reduced it. Despite its excitatory action, 5-HT did not facilitate visual responses. Spontaneous changes in ongoing activity were found to affect the visual response amplitude in the same way. Noradrenaline, the alpha 1-agonist PHE and the beta-agonist ISO exerted a weak depressant action on high-frequency maintained activity, but during low-frequency single spike activity and/or burst activity a facilitatory effect was evident, which prevented the generation of burst discharges and slightly increased single spike firing. Visually evoked activity was little affected, but signal-to-noise ratio changed with changes in ongoing activity. The alpha 2-agonist CLO clearly attenuated both spontaneous activity and visual responses. We suggest that, in addition to direct effects of ACH and 5-HT on geniculate relay cells, the balance between the opposite actions of ACH and 5-HT on PGN cells determines the mode of operation in the recurrent inhibitory circuit: either a global, tonic inhibition of relay cells during a dominating 5-HT influence or a less tonic but phasic inhibition during increased activity in the cholinergic system.

Acetylcholine

Immunohistochemical staining for glial fibrillary acidic protein (GFAP) after deafferentation or ischemic infarction in rat visual system: features of reactive and damaged astrocytes.

Immunohistochemical staining for glial fibrillary acidic protein (GFAP) is standard for visualization of reactive astrocytes in tissue sections, whereas various forms of astrocytic damage remain to be described in detail. In this study we tested differences in GFAP labeling in reactive astrocytes and in glial cells damaged by ischemia and edema. Studies were performed in the anatomically well defined visual system of rat. Basic staining patterns for GFAP were established in subcortical visual nuclei and visual cortex. In the first model, deafferentation of visual centers was performed by unilateral optic nerve lesion, and characteristic changes of GFAP labeling in reactive astrocytes were studied at 0.5, 1, 1.5, 2, 4, 8 and 21 days after lesion. Initial changes were seen in the deafferented superior colliculus at 1 day after deafferentation with a diffuse increase and stellate types of reactive cells formed at 2-8 days. In the second model, small ischemic infarcts were produced in the visual cortex of rats using the method of photochemically-induced thrombosis. GFAP labeling with a polyclonal antiserum was massively enhanced in the infarct at 4 hr. Characteristic morphological changes in damaged astrocytes were seen which were also identified in experiments with simulated global ischemia. In the surround of the infarct, swelling of astrocytes also caused increased labeling. At 3-4 days infarction typical reactive astrocytes surrounded the lesioned area. In conclusion, these immunohistochemical studies on GFAP in rat visual system allow for the following classifications. (a) Normal astrocytes vary in labeling at different anatomical localizations. (b) Reactive astrocytes show enhanced labeling and larger cell-size within an interval of 1-2 days after lesion. (c) Astrocytes damaged by ischemia reveal increased labeling of disintegrating cellular elements within hours after a lesion. (d) Swollen astrocytes undergo enhanced labeling in areas with vasogenic edema.

Animals

Noradrenergic modulation of retinogeniculate transmission in the cat.

1. Relay neurones were extracellularly recorded from the A-layers of the dorsal lateral geniculate nucleus (dLGN) of the anaesthetized cat. The noradrenergic influence on retinogeniculate transmission was investigated through microiontopheretic techniques in a total of 140 dLGN relay cells using three experimental approaches: (i) the effects of agonists for alpha 1-, alpha 2- and beta-adrenoceptors were separately analysed; (ii) the noradrenergic influence was related to the global state of activity of the relay neurones, which was associated with discrete patterns of the electroencephalogram (EEG); (iii) distinct phases of visual responses evoked from the area of the retinal receptive field, and of binocular and lateral inhibitory responses, were evaluated before, during and after the action of noradrenergic agonists. 2. The spontaneous generation of high-frequency bursts of spikes in dLGN relay neurones, associated with periods of highly synchronized, delta-like patterns of the EEG, was selectively suppressed by the beta-adrenoceptor agonist isoprenaline or the alpha 1- adrenoceptor agonist phenylephrine. Single action potentials, occurring at a low frequency between bursts, were significantly less affected. Depending upon the ejection level of the adrenoceptor agonists, burst activity was suppressed by 23-73%, compared with a reduction in single spike firing in the range 7-24%. The suppression of burst firing occurred in all functional types of dLGN relay neurones (X, Y; on, off), enhanced burst activity was observed in less than 1% of the cells. 3. On-going tonic sequences of action potentials (around 15 Hz), occurring during periods of EEG activity characterized by lower amplitudes and higher frequencies, were separately affected by adrenoceptor agonists. Isoprenaline had no significant effect, phenylephrine induced a global reduction of spike firing with no obvious relation to the ejection level, and the alpha 2-adrenoceptor agonist clonidine inhibited action potential generation in a near dose-dependent manner. 4. Visual response properties were investigated during periods of less synchronized states of EEG activity. Responses to visual stimulation of the retinal receptive field centre were not significantly influenced by isoprenaline, while phenylephrine or clonidine attenuated the phasic and the tonic response component in all functional types of relay neurones and independent of the stimulus contrast being used. At low ejection levels, slight facilitatory effects were observed with isoprenaline (65% of neurones that were tested) or phenylephrine (15%). The inhibitory influence of the antagonistic surround area of the receptive field appeared unaltered during action of isoprenaline or phenylephrine.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials

EEG-dependent modulation of response dynamics of cat dLGN relay cells and the contribution of corticogeniculate feedback.

Single unit recordings were made extracellularly from the dorsal lateral geniculate nucleus (dLGN) in the anaesthetized and paralysed cat. The impulse rates of phasic (peak) and tonic components of visual responses to stimulation of the receptive field center by a flashing spot were determined during different states of the EEG, during local cortical cooling and during micro-iontophoretic application of the excitatory amino acid receptor agonists, quisqualate (QUIS) and N-methyl-D-aspartate (NMDA). Typically, visual responses were phasic during low frequency/high amplitude EEG patterns, resembling slow wave sleep (SWS). During high frequency EEG patterns (non-SWS) visual responses of X- and Y-cells exhibited a prominent tonic response component. This tonic component could be clearly reduced during ipsilateral cortical cooling in the non-SWS state. QUIS or NMDA, applied in order to mimic corticofugal activity, augmented the tonic response component, most efficiently during SWS EEG. The effects did not differ significantly for X- and Y-cells. During non-SWS EEG Y-cells exhibited a tonic response component similar to X-cells, but because of their higher peak rates the responses of Y-cells were on average more phasic than those of X-cells. Our results indicate that state-dependent changes in CNS activity modulate dL'GN responsiveness in part via the corticogeniculate feedback, and that predominantly the tonic response component is modulated.

Animals

Immunohistochemical changes of neuronal calcium-binding proteins parvalbumin and calbindin-D-28k following unilateral deafferentation in the rat visual system.

The neuron-specific calcium-binding proteins, parvalbumin and calbindin-D-28k, were studied in the subcortical visual system of normal and unilaterally deafferented albino rats. Immunohistochemistry with monoclonal antibodies was used on vibratome sections through optic tract (OT), dorsal lateral geniculate nucleus (dLGN), olivary pretectal nucleus (OPN), and superior colliculus (SC). In controls, OT stained strongly for parvalbumin and weakly for calbindin-D-28k. The dLGN contained a plexus of parvalbumin-positive fibers. In dLGN, calbindin-D-28k-antibodies showed strong labeling of some neurons with long dendrites and weak staining of the cytoplasm in other neurons. In OPN, parvalbumin stained a ring of neurons and terminals in the shell region, whereas calbindin-D-28k was contained in medial cell populations. In SC, parvalbumin was contained in fibers, terminals, and neurons throughout the visual layer. Calbindin-D-28k showed a laminar distribution of neurons with a predominance in deep portions of superficial grey matter and in ventral portions of stratum opticum. Following unilateral deafferentation induced by optic nerve section, retinal axons showed immunohistochemical changes related to Wallerian degeneration and target neurons reacted by changes of calcium-binding proteins. Parvalbumin and calbindin-D-28k immunostaining decreased during Wallerian degeneration of OT. In the deafferented dLGN, immunohistochemical labeling for calbindin-D-28k declined in strongly stained neurons from 4 to 21 days after lesion. Measurement of dendritic length per number of cells or per area of dLGN showed a significant decline for the contralateral side at 4, 8, and 21 days (ANOVA, P less than 0.05). In deafferented OPN, terminal-like staining for parvalbumin decreased and neuronal labeling was enhanced. In deafferented SC, the neuronal and dendritic staining for parvalbumin increased beginning from Day 1 on and persisting at Day 21, whereas fibers and terminal-like elements decreased in staining. Measurement of parvalbumin-positive neurons per area of SC showed a significant increase of labeling in the contralateral side from Day 1 to Day 21 (ANOVA, P less than 0.05). These studies show that cellular responses to deafferentation of visual neurons involve a regulation of calcium-binding proteins. The decline in staining for calbindin-D-28k in dLGN may relate to reduced retinal afferent activity. The progressive cellular changes in parvalbumin staining may be related to unmasking of intrinsic neurons after removal of parvalbumin-containing, afferent fibers and terminals. Additionally, the changes of parvalbumin labeling in SC neurons may reflect a plastic reorganization of local circuits known to occur in rat SC in response to deafferentation.

Afferent Pathways

Cellular organization of reciprocal patchy networks in layer III of cat visual cortex (area 17).

There is no direct information available concerning the exact spatial characteristics of long-range axons and their relationship with the patchy phenomena observed after extracellular injection of retrograde tracers. In the present study, using the recently introduced neuronal tracer biocytin, we demonstrate by detailed three-dimensional reconstruction of 10 pyramidal cells in layer III, that their clustered axonal terminals form a specific patchy network in layers II and III. The reconstructed network occupied an area of 6.5 x 3.5 mm parallel to the cortical surface elongated in an anteroposterior direction. The average centre-to-centre distance between patches within the network was 1.1 mm. On average, the axonal field of each of the 10 pyramidal cells contained a total of 417 boutons at four to eight distinct sites (patches), and in each patch, an average of 79 boutons was provided by the same cell. The identified connections between the patches were predominantly reciprocal. Detailed analyses have shown that many pyramidal cells of the network are directly interconnected so that each of them can receive one to four, chiefly axospinous, contacts onto the distal segment of its apical and basal dendrites from the axon of another pyramidal cell belonging to a different patch labelled from the same injection site. We hypothesize that the possible functional role of the network is to link remote sites with similar physiological characteristics, such as orientation preference, supporting the model of Mitchison and Crick [(1982) Proc. natn. Acad. Sci. U.S.A. 79, 3661-3665].

Animals