Bibliothek

Notizen: Summary The responses to moving stimuli of single cells in the parvo- and magnocellular layers (PCL and MCL) of the macaque lateral geniculate nucleus (LGN) have been studied. PCL cells respond with a monophasic increase or decrease in firing when a bar passes across the receptive field, according to the wavelength composition of the stimulus. MCL cells respond with a biphasic sequence of excitation and suppression or vice versa dependent on whether a cell is on-centre or off-centre and on stimulus contrast direction. With large stimuli, PCL cells respond as long as the stimulus covers the receptive field while MCL cells respond only at the contrast borders. MCL cell responses are maximal with bars just long enough to cover the field centre, while PCL cell responses show a variable relation with bar length, depending on stimulus wavelength and receptive field structure. PCL cells show broad velocity tuning while at least some MCL cells were more sharply tuned. Many cells in the macaque LGN show weak orientation or direction preference.

Notizen: Summary The receptive fields of LGN cells were investigated with stationary light and dark spot and annulus stimuli. Stimulus size and background intensity were varied while stimulus/background contrast was kept constant. The speed of dark adaptation varied considerably from cell to cell. Dark adaptation made responses more sustained in all neurones and eliminated the oscillatory on-responses evoked under some conditions in the light-adapted cells. Dark adaptation led also to a disappearance of early phasic inhibition in on-responses, and increased response rise time and latency. The power of surround responses to inhibit centre responses decreased slightly at low levels of light adaptation in LGN cells but much less than in retinal ganglion cells. Some other traces of changing retinal surround effects also appeared in the LGN on dark adaptation. For example, the functional size of receptive fields increased at low levels of illuminance as has been observed in retinal ganglion cells and the receptive fields as estimated from response peaks were larger than those estimated from sustained components.

Notizen: Summary Steadily illuminated surrounds, remote from the receptive field centre, are shown to affect the responses of primate visual cells. Intensity-response curves of cells of the macaque lateral geniculate nucleus were measured using a successive contrast paradigm where chromatic or achromatic stimuli were presented in alternation with a white adaptation field of constant luminance. Adding white surround annuli around stimuli and adaptation field shifted the intensity-response curves to higher intensity ranges. Since response curves can be non-monotonic, this remote surround effect can result in an increase or decrease in responsiveness (facilitation or suppression) dependent on stimulus intensity. Steady surrounds, remote from the receptive field centre, thus control cell sensitivity and responses by means of simultaneous contrast.

Notizen: 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.

Notizen: 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.

Notizen: 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.

Notizen: Summary The responses of neurones in laminae A and A1 of the cat lateral geniculate nucleus to moving stimuli were investigated at different background luminances. Moving bright slits, dark bars and edges were employed; the contrast of stimuli against the background was held constant. Background intensities varied from 10−3 to 102 td. Responses as stimuli passed across the centres of LGN receptive fields became stronger with increasing levels of light adaptation up to 10−1–101 td and then remained constant. Responses as stimuli passed through surround regions altered qualitatively with adaptation level, generally increasing in strength and complexity with background luminance. As a bright slit for on-centre cells or dark bar for off-centre cells left the surround, in almost all units a strong secondary peak could be elicited by an appropriate selection of the adaptation conditions. Many features of the responses to moving stimuli could not be predicted from the responses to stationary stimuli under different adaptation conditions described in the previous paper.