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Notes: Summary The afferent and efferent connections of the prelunate visual association area V4 of macaque monkeys were investigated by means of the horseradish peroxidase (HRP) method. The specific thalamic afferents from the dorsolateral segment of the medial pulvinar and the lateral segment of the inferior pulvinar were topographically organized. A band of cells was labelled in the intralaminar nuclei (nucl. centr. med. and lat., reaching into LD and the most dorsal part of VL), and a few cells in the interlaminar layers of the lateral geniculate body. Other diencephalic afferents included the claustrum, the nucleus basalis Meynert and the pars compacta of the substantia nigra. Ipsilateral cortical areas which projected into V4 included area 18 (V2), the inferior parietal cortex, the anterior and posterior parts of the superior temporal sulcus, the frontal eye fields and the temporo-basal association cortex on the lateral half of the parahippocampal gyrus and around the occipito-temporal sulcus. In the contralateral cortex, discontinuous regions in areas V4 and V5 on the prelunate gyrus and some cells at the 17/18-border were labelled. All regions in which labelled cells were found and, in addition a restricted region in the dorsal cap of the head and the tail of the caudate nucleus showed fibre and terminal labelling. In addition mesencephalic afferents and efferents were identified but not investigated in detail. An attempt to estimate the quantitative contribution of the various afferent systems to the prelunate cortex was made by counting the labelled cells in the different areas. The afferent and efferent organization of the prelunate visual association area indicates that it is incorporated in a network of cortical and subcortical regions involved in various aspects of visual behavior.

Notes: Summary In the common marmoset (Callithrix jacchus), the cortical projection from the pulvinar and other diencephalic structures into the striate and prestriate cortex was investigated with various fluorescent retrograde tracers. Single cortical injections as well as multiple injections at distances of 1–2 mm with one tracer into an extended but coherent cortical region were applied. Fields with multiple injections were placed so that they touched each other (minimal distances 2 to 3 mm). Retrogradely labelled cells in the LGN and/or the pulvinar were arranged in coherent columns, volumes or slabs, but cell volumes resulting from neighbouring cortical injections overlapped at their border (for details of the thalamo-cortical topography see the companion paper Dick et al. (1991)). Double labelled cells (dl) were only found in the zones of overlap of the cell volumes labelled by the respective tracers. The relative number of dl-cells in these overlap zones was 6.2 ± 3.1%. The dl-frequency was the same in the various nuclei of the pulvinar and the LGN. In the main layers of LGN, dl-cells were found only in the overlap zone of two injection fields into area 17, but a few dl-cells were found in interlaminar cells after injections into area 17 and 18. Maximal cortical distances between injection fields which produced dl in the pulvinar, were 3 to exceptionally 4 mm but dl was highest at injection distances ≤2.5 mm and decreased sharply at wider distances. Such overlap zones were concerned with identical or overlapping regions of visual field representation in the cortex and probably also in the pulvinar. Although in individual experiments up to four different tracers were injected into different striate/prestriate regions, often embracing the same visual field representation, individual cells in the pulvinar showed dl from maximally only two tracers injected into neighbouring cortical regions. We conclude that dl in the posterior thalamic projection nuclei is determined essentially by cortical distance and thus reflects the local domain of branching of thalamo-cortical afferents. Pruning of such branches during development may further restrict bifurcating axons to identical visual field representations, but representation of identical visual field regions in different visual areas is not, per se, a sufficient condition for dl. It is not found if such regions are further apart from each other than the typical local domain of 2–3 mm, exceptionally up to 4 mm in one experiment after injections into area 17 and MT. Dl in the intralaminar nucleus CeL (5.0 ± 4.6%), the claustrum (5.4 ± 3.6%) and in the amygdala (5.7 ± 1.9%) was of the same order as in the pulvinar and LGN. In the hypothalamus around 10% and in the Nucleus basalis Meynert 15.8% of the cells labelled by visual cortical injections were double labelled. In all these extrathalamic regions dl was also restricted to overlap zones, but overlap of labelled fields in these nuclei was much wider and included the whole striate/prestriate cortex except for some topographical separation of striate and prestriate projection zones in the claustrum. Only in the Nucl. basalis Meynert and the hypothalamus some cells were labelled by three tracers.

Notes: Summary The distribution of mRNA expression for three types of voltage gated neuronal sodium-channels was studied in the rat brain at different developmental stages (embryonal day E18, postnatal day P5 and adult). With the in-situ hybridization technique, using synthetic DNA-oligomer probes, pronounced regional and temporal variations in the expression levels of the different channel subtypes could be detected. In comparison with types I and III, sodium channel II mRNA was the most abundant subtype at all developmental stages. Maximal expression of sodium channel II mRNA was seen at P5 in virtually all parts of the grey matter, except for the cerebellum. In adult rat brain in contrast, sodium channel II mRNA levels were maximal in the granular layer of the cerebellum, whereas in all other regions expression had decreased to roughly 50% of postnatal levels. Na channel I expression was virtually absent at E18 and showed highest levels at P5, with maxima in the caudate nucleus and hippocampus. In the adult brain, expression of Na-channel I was nearly absent in the neocortex, but well detectable in the cerebellum and, at lower levels in the striatum and thalamus. Sodium channel III was mainly expressed at the embryonal stage and showed a decrease to very low levels with little regional preferences in the adult.

Notes: Summary Over 300 single units from the visual cortex (within and around the projection of the central area) were recorded from awake and non-paralyzed cats (chronic preparation). Spontaneous activity of 25% of the neurons was below 3/sec, that of 75% above 3/sec (mean 7.65 spikes/sec). Diffuse illumination had only little influence, but nearly all neurons responded to stimulation with some sort of visual contrast. This would be either an irregularly moved shadow on the screen with irregular boundaries (e. g. a hand with moving fingers), a dark stripe moving in a certain direction, stationary parallel gratings with a certain orientation, or saccadic eye movements across a checkerboard. Although some neurons responding to one stimulus type could also be responsive to other stimuli, the majority of units only responded to one stimulus type. The responses to stationary gratings (alternating parallel dark and bright stripes) and to moving dark stripes are described in detail. Responses to stationary gratings showed no adaptation. The orientation of the grating stripes was critical for each neuron, the optimal and minimal response orientation were separated by about 90°. For movement sensitive neurons, the direction of the movement was critical. Most neurons had only one, some had two preferred directions separated by 180°. No statistically significant predominance of certain orientation or direction preferences was found. The preferred target velocity of movement sensitive neurons was between 10 and 60°/sec, above 80–100°/sec only occasional or no responses could be elicited. Neurons which responded to saccadic eye movements (above 300°/sec) in the presence of a checker board, usually did not respond to slower target movements below 100°/sec. The results support the view that the visual system has different channels for the perception of moving and of stationary objects.

Notes: Summary The colour of an object is changed by surround colours so that the perceived colour is shifted in a direction complementary to the surround colour. To investigate the physiological mechanism underlying this phenomenon, we recorded from 260 neurons in the parvo-cellular lateral geniculate nucleus (P-LGN) of anaesthetized monkeys (Macaca fascicularis), and measured their responses to 1.0–2.0° diameter spots of equiluminant light of various spectral composition, centered over their receptive field (spectral response function, SRF). Five classes of colour opponent neurons and two groups of light inhibited cells were distinguished following the classification proposed by Creutzfeldt et al. (1979). In each cell we repeated the SRF measurement while an outer surround (inner diameter 5°, outer diameter 20°) was continuously illuminated with blue (452 nm) or red (664 nm) light of the same luminance as the center spots. The 1.0–1.5° gap between the center and the surround was illuminated with a dim white background light (0.5–1cd/m2). During blue surround illumination, neurons with an excitatory input from S-or M-cones (narrowand wide-band/short-wavelength sensitive cells, NSand WS-cells, respectively) showed a strong attenuation of responses to blue and green center spots, while their maintained discharge rate (MDR) increased. During red surround illumination the on-minus-off-responses of NS- and WS-cells showed a clear increment. L-cone excited WL-cells (wide-band/long-wavelength sensitive) showed a decrement of on-responses to red, yellow and green center spots during red surround illumination and, in the majority, also an increment of MDR. The response attenuation of narrow-band/long-wavelength sensitive (NL)-cellls was more variable, but their on-minus-off-responses were also clearly reduced in the average during red surrounds. Blue surround illumination affected WL-cell responses little and less consistently than those of NL-cells, but often broadened the SRF also in the WL-cells towards shorter wavelengths. The M-cone excited and S-cone suppressed WM-cells were strongly suppressed by blue but only little affected by red surround illumination. The changes of spectral responsiveness came out clearly in the group averages of the different cell classes, but snowed some variation between individual cells in each group. The zero-crossing wavelengths derived from on-minus-off-responses were also characteristically shifted towards wavelengths complementary to those of the surround. The direction of changes of spectral responsiveness of P-LGN-cells are thus consistent with psychophysical colour contrast and colour induction effects which imply that light of one spectral region in the surround reduces the contribution of light from that same spectral region in the (broad band or composite) object colour. Surrounds of any colour also decrease the brightness of a central coloured or achromatic light (darkness induction). We calculated the population response of P-LGN-units by summing the activity of all WS-, WM- and WL-cells and subtracting that of all NS- and NL-cells. The SRF of this population response closely resembled the spectral brightness function for equiluminous lights rather than the photopic luminosity function. With red or blue surrounds, this population SRF was lowered nearly parallel across the whole spectrum to about 0.7 of the amplitude of the control. In a psychophysical test on 4 observers we estimated the darkness induction of an equiluminous surround in a stimulus arrangement identical to the neurophysiological experiment, and found a brightness reduction for white, blue, green and red center stimuli to 0.5–0.7 of the brightness values without surround. This indicates that the neurophysiological results may be directly related to perception, and that P-LGN-cells not only signal for chroma but also for brightness, but in different combinations. The results indicate that both an additive (direct excitation or suppression of activity) and a multiplicative mechanism (change of gain control) must be involved in brightness and colour contrast perception. As mechanisms for the surround effects horizontal cell interactions appear not to be sufficient, and a direct adaptive effect on receptors feeding positive or negative (opponent) signals into the ganglion cells receptive fields by straylight from the surround must be seriously considered. This will be examined in the following companion paper. The results indicate that changes of spectral and brightness responses in a colour contrast situation sufficient to explain corresponding changes in perception are found already in geniculate neurons and their retinal afferents. This applies to mechanisms for colour constancy as well in as much as they are related to colour contrast.

Notes: Summary We report on experiments which were undertaken in an attempt to clarify mechanisms underlying the contrast effects of chromatic surround illumination on spectral responsiveness of cells in the parvocellular layers of the LGN (P-LGN-cells), that had been demonstrated under standard conditions in the preceding companion paper. The experiments were done in anesthetized macaques (Macaca fascicularis). In some neurons, S-potentials were recorded together with the post-synaptic action potentials, and all effects seen in P-LGN-cells were present already in their retinal afferents indicating their retinal origine. The responsiveness of the cells for center stimuli of different wavelengths and during illumination of the receptive field center or the outer surround was determined. Continuous outer surround illumination alered maintained dicharge rate (MDR), sensitivity and gain of P-LGN and retinal ganglion cells in the same way and empirically not distinguishable from direct illumination of the receptive field. Responses to surround flashes showed the same dependence on spectral composition as those to center flashes. Adaptation and excitation caused by outer surround illumination (inner diameter 5°, outer diameter 20°) were, in the average, ten times weaker than those exerted by light of the same spectral composition shone directly into the receptive field. Surround effects decreased proportional to r-2. Excitation by outer surround flashes was reduced by adaptation of the receptive field center in the same manner as responses to center flashes. The findings indicate that outer surround light has a direct excitatory and adaptive effect on the excitatory or inhibitory cones feeding into the receptive field. This indicates that straylight from the surround into the center could be responsible for the adaptive and excitatory effects of surround illumination. The straylight fraction from the remote surround into the receptive field must be higher, however, than that estimated from the psychophysically determined point spread function. It comes closer to earlier direct straylight measurements in excised eyes, but may be enhanced by chromatic aberration. If a surround of excitatory colour is flashed simultaneously with an excitatory center stimulus, additivity of center and surround excitation is observed only at low center intensities, while at higher center intensities the gain for center excitation is reduced similar to adaptive gain control. This could be explained by lateral interaction through horizontal connections in the retina, which decays within seconds, while adaptation of the cones feeding into the receptive field center is fully effective only after about 3 s. Our findings therefore suggest a two stage model for surround effects, a fast one mediated through horizontal connections controlling the gain of receptorbipolar transmission and a slow one through adaptation by straylight and controlling receptor gain. The fast process is receptor unspecific, i.e. pooling activity from all receptor types, while the second one is receptor specific. During real seeing both processes are simultaneous and complement each other because of continuous eye movements. Perceptual darkness and colour induction by remote surrounds are consistent with this model, which can also be applied to colour constancy. WM-cells (Yellow-minus-blue) show peculiar properties during surround or center illumination with blue light, suggestng an opponent mechanism different from that of other P-LGN-cells.

Notes: Summary In the lateral geniculate body (LGB), intra- and quasi-intracellular records were done. With small light stimuli shone into different parts of the receptive field, EPSPs and IPSPs could be elicited. Stimulation of the exact center of an on-center cell produced a pure excitatory response, that of an off-center neurone pure inhibition. This response lasted throughout the stimulus. At light off, inhibition was elicited in on-center cells and excitation in off-center cells. A stimulus in the field periphery produced a mixed response with a small and short excitation followed by large inhibition in on-center cells, and a short inhibition followed by postsynaptic depolarization in off-center cells. At light off, on-center cells showed depolarization after a short polarizing phase, and off-center cells a broad polarization which interrupted the initial small excitation. The latencies of both the excitatory and inhibitory center responses at light on and off characteristic for the two types of neurones, were 20–30 msec shorter than the reversed responses elicited by stimulation of the receptive field surround. The findings are compatible with a model in which each geniculate on-center cell gets its major excitatory input from one optic tract on-center fibre and inhibitory input from several off-center fibres with nearby receptive fields. An off-center LGB-cell receives its main excitation essentially from one offcenter fibre and inhibition from several on-center cells. The responses to moving stimuli also agreed with this model. The presence of recurrent inhibition within the LGB could be confirmed by electrical stimulation. But it could not be decided whether the reciprocal inhibition of on- and off-center cells was due to forward or backward inhibition. The spontaneous activity of on- and off-center cells which were simultaneously recorded with one electrode, showed a mutual inhibition 6–8 msec after one cell had fired. Anatomical data relevant to the model are discussed and some functional implications are suggested.

Notes: Summary The effect of monocular and binocular stimulation on cortical neurons of area 17 was investigated in awake unparalyzed cats with painless head fixation. Two types of stimuli were applied: Stationary gratings of variable orientations, and a 3° wide dark stripe at different orientation and moving in different directions. All neurons which were excited from both eyes showed qualitatively similar input properties (orientation specificity, movement and/or direction sensitivity). Quantitatively, the input from both eyes was either equal or dominant from one eye. Contralateral dominance was found 5 times more frequently than ipsilateral dominance. Various types of binocular interaction were found. Some neurons showed an excitatory response from one eye and inhibitory response from the other (inhibition, 14% of our units), and others showed a response during binocular stimulation which was equal to the sum of the monocular responses (summation, 18%), larger (facilitation, 43%) or smaller (occlusion, 14%) than the sum of the two monocular responses. A few units with binocular responses did not respond to monocular stimulation of one or both eyes. The results are compared with those found by other authors in paralyzed and anesthetized animals, and current theories of neuronal mechanisms of binocular vision are discussed in the context of our findings.

Notes: Summary 1. Responses of cortical cells from the foveal and perifoveal visual field representation in area 17 to moving contrasts were analyzed with intracellular records in anesthetized cats. These intracellularly recorded responses were normal in so far as the cells showed typical orientation/direction sensitivity and only short phasic or no responses to diffuse illumination. 2. With slowly moving bright or dark bars, two types of responses were seen: those with a small excitatory peak and those with a wider excitatory peak. Inhibitory regions outside the excitatory peak were only seen in cells with a small excitatory area. Only very few cells showed inhibitory “flanks” preceding and following the excitation; often inhibition followed the excitation in both the forward and backward direction; sometimes it preceded it in both directions. The inhibition outside the excitatory zone practically always had “dynamic” properties, i.e. was smaller or larger in the two opposite directions of movements. 3. All cells showed strong inhibition (IPSP's) mixed with excitation while the stimulus moved over the excitatory response field. The degree of inhibition was clearly sensitive to the direction of movement (forward or backward) of an optimally oriented moving stimulus, and could also be different at different orientation/ directions. However, the orientation dependence of intracortical inhibition was often less clear than the differences found between the two opposite directions of an optimally oriented stimulus. Inhibition was more marked during binocular than during monocular stimulation. 4. The excitatory areas of cortical cells were mostly slightly elongated, but not systematically along the axis of optimal orientation. The diameters of the excitatory fields were similar along the optimal and the non-optimal orientation axes (mean 1.9±0.78 vs. 2.2±0.92°). 5. It is proposed that the orientation/direction sensitivity of cortical cells is a function of intracortical inhibitory connections with direction/orientation sensitivity rather than only due to the spatial arrangement of excitatory and inhibitory on- or off-center fields. A hypothetical retino-cortical projection map is proposed and it is assumed that direction/orientation sensitive intracortical inhibition is essential for the functional properties of cortical neurones.

Notes: Summary In eleven hemispheres of nine marmoset monkeys (Callithrix jacchus), we have investigated the thalamo-cortical organization of the projections from the pulvinar to the striate and prestriate cortex. In each experiment, single or multiple injections of various retrograde fluorescent tracers were injected into adjacent regions or areas. In two experiments, horseradish peroxidase (HRP) was injected into the lateral geniculate nucleus (LGN) and the lateral pulvinar, respectively. The results show that the thalamo-cortical projection from LGN to striate cortex and from pulvinar to the prestriate cortex are similarly organized, but the geniculo-striate projection is more precise than the pulvinar-prestriate projection. The pulvinar-prestriate projection is topographically organized and preserves topological neighbourhood relations. Projection zones to the various visual areas are concentrically wrapped around each other. The projection zone to area 18 constitutes a central core region. It begins ventro-laterally in PuL where the pulvinar is in contact with the LGN. This contact zone we called the hilus region of the pulvinar. The area 18-projection zone stretches as a central cone into the posterior pulvinar through PuL and into PuM. It is surrounded by the projection zone to the posterior belt of area 19 and this in turn is surrounded by the projection zone to the anterior belt of area 19. The projection zones to area 19 are then surrounded medially and dorsally by zones projectiong to the temporal and parietal association cortex, respectively. The projection zone to area MT is located medio-ventrally in the posterior pulvinar (PuIP and surrounding nuclei) and coincides with a densely myelinated region. Area 17 also receives input from the pulvinar but probably predominantly in the region of the central visual field. The pulvinar zone projecting to area 17 is located ventrolaterally from the central core region projecting to area 18 and is contiguous laterally with the LGN. If the positions of the vertical and the horizontal meridian in the pulvinar correspond to those in the respective cortical projection zones, a second order visual field representation such as found in area 18, with the horizontal meridian split at an excentricity of about 7–10°, can also be recognized in the pulvinar.