1. It has recently been shown that the X- and Y-cell classes in the A-layers of the cat lateral geniculate nucleus (LGN) are divisible into lagged and nonlagged types. We have characterized the visual response properties of 153 cells in the A-layers to 1) reveal response features that are relevant to the X/Y and lagged/nonlagged classification schemes, and 2) provide a systematic description of the properties of lagged and nonlagged cells as a basis for understanding mechanisms that affect these two groups. Responses to flashing spots and drifting gratings were measured as the contrast and spatial and temporal modulation were varied. 2. X- and Y-cells were readily distinguished by their spatial tuning. Y-cells had much lower preferred spatial frequencies and spatial resolution than X-cells. Within each functional class (X or Y), however, lagged and nonlagged cells were similar in their spatial response properties. Thus the lagged/nonlagged distinction is not one related to the spatial domain. 3. In the temporal domain X- and Y-cells showed little difference in temporal tuning, whereas lagged and nonlagged cells showed distinctive response properties. The temporal tuning functions of lagged cells were slightly shifted toward lower frequencies with optimal temporal frequencies of lagged X-cells averaging an octave lower than those of nonlagged X-cells. Temporal resolution was much lower in lagged X- and Y-cells than in their nonlagged counterparts. 4. The most dramatic differences between lagged and nonlagged cells appeared in the timing of their responses, as measured by the phase of the response relative to the sinusoidal luminance modulation of a spot centered in the receptive field. Response phase varied approximately linearly with temporal frequency. The slope of the phase versus frequency line is a measure of total integration time, which we refer to as visual latency. Lagged cells has much longer latencies than nonlagged cells. 5. The intercept of the phase versus frequency line is a measure of when in the stimulus cycle the cell responds: we refer to this as the intrinsic or absolute phase of the cell. This measure of response timing not only distinguished lagged and nonlagged cells well but also covaried with the sustained or transient nature of cells' responses to flashed stimuli.(ABSTRACT TRUNCATED AT 400 WORDS)
Horseradish peroxidase was injected intracellularly into single, physiologically-identified X- and Y-cell geniculocortical axons projecting to area 17 of the cat. This injection anterogradely labeled the axon terminal fields in cortex and retrogradely labeled the somata of these same axons in laminae A and A1 of the lateral geniculate nucleus (LGN). The laminar projections of 21 X- and 15 Y-cell axons were analyzed. For these, the laminar terminations of ten X- and seven Y-cell axons were also related to their cells' positions in the A-laminae. The terminal fields of X- and Y-cell axons overlapped substantially in layers IV and VI of area 17. Some X-cells terminated mainly in IVb, others mainly in IVa, and still others throughout IVa and IVb. The latter two groups also projected up to 100 micron into lower layer III. Y-cells terminated primarily in layer IVa and projected up to 200 microns into lower layer III. Some also arborized throughout the depth of layer IVb. Both X- and Y-cell axons terminated throughout the depth of layer VI, although more so in the upper half. We found no relationship between the diameter of the parent axon and its sublaminar projection within layer IV. Within layer IV, X-cell axons generally terminated within a single, continuous clump and had surface areas of 0.6 to 0.9 mm2. Axons of Y-cells often terminated in two to three separate clumps, separated by terminal free gaps 400 to 600 micron wide. Their total surface areas, including gaps, were 1.0 to 1.8 mm2, roughly 1.6 times the surface areas of X-cell axons. Despite considerable overlap, Y-cell arbors contained significantly more boutons than did X-cell arbors. The sublaminar projections of the X- and Y-cell axons within layer IV reflected the locations of the cells' somata within the depth of the A-laminae. X-cells located in the dorsal or ventral thirds of the depths of the laminae projected mainly to layer IVa or throughout layer IV in cortex. Those located in the central thirds projected mainly to layer IVb. Y-cells showed a similar positional relationship, but they appeared to follow different rules. Y-cells in the outer thirds of the A-laminae projected mainly to layer IVa; those in the central thirds, in addition, expanded their projections to include layer IVb. In general, larger sized somata in the LGN gave rise to more widely spreading terminal arbors and greater numbers of boutons in cortex than did smaller somata.(ABSTRACT TRUNCATED AT 400 WORDS)
1. The lateral geniculate nucleus is the primary thalamic relay through which retinal signals pass en route to cortex. This relay is gated and can be suppressed by activity among local inhibitory neurons that use gamma-aminobutyric acid (GABA) as a neurotransmitter. In the cat, a major source of this GABAergic inhibition seems to arise from cells of the perigeniculate nucleus, which lies just dorsal to the A-laminae of the lateral geniculate nucleus. However, the morphological characteristics of perigeniculate cells, and particularly the projection patterns of their axons, have never been fully characterized. We thus examined the morphology of these cells: individually by intracellular injection of horseradish peroxidase (HRP) and en masse with the anterograde tracer Phaseolus vulgaris leucoagglutinin (PHAL). 2. We recorded from 12 perigeniculate cells that we impaled and successfully labeled with HRP. These cells exhibited response properties generally consistent with those described previously. They had long response latencies to stimulation of the optic chiasm and relatively large, often diffuse, receptive fields. The visually evoked responses of most of the cells were dominated by one eye. Compared with cells of the lateral geniculate nucleus, perigeniculate cells had large somata (517 +/- 136 microns 2 in cross-sectional area, mean +/- SD), which were fusiform or multipolar in shape, and dendritic arbors that extended a considerable distance (1,095 +/- 167 microns) parallel to the border between the perigeniculate and lateral geniculate nuclei. Terminal arbors of some dendrites were quite complex and beaded. 3. The axons of six perigeniculate cells were labeled sufficiently well to trace and reconstruct over a considerable distance. Each of these axons formed branches that descended to innervate the lateral geniculate nucleus, and this geniculate innervation was exclusively limited to the A-laminae. Terminal boutons within the A-laminae were nearly all en passant, which gave the axons a beaded appearance. Furthermore, branches of five of these six axons provided local innervation of the perigeniculate nucleus, generally within each labeled cell's own dendritic arbor. Three of the cells also exhibited an axon branch that extended medially and caudally away from the soma, but we were unable to trace these axon branches to their targets. 4. Within the lateral geniculate nucleus, each arbor of perigeniculate axons derived from two main components. One was a narrow, sparse medial component that innervated laminae A and A1.(ABSTRACT TRUNCATED AT 400 WORDS)
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