The recent appreciation of the fact that the pulvinar and lateral posterior (LP) nuclei receive two distinct types of cortical input has sparked renewed interest in this region of the thalamus. A key question is whether the primary or "driving" inputs to the pulvinar/LP complex originate in cortical or subcortical areas. To begin to address this issue, we examined the synaptic targets of tectothalamic terminals within the LP nucleus. Tectothalamic terminals were labeled using the anterograde transport of biotinylated dextran amine (BDA) or Phaselous leucoagglutinin placed in the superior colliculus or using immunocytochemical staining for substance P, a neurotransmitter found to be used by the tectothalamic pathway (Hutsler and Chalupa [ 1991] J. Comp. Neurol. 312:379-390). Our results suggest that most tectothalamic terminals are large and occupy a proximal position on the dendritic arbor of LP relay cells. In the medial LP, tectothalamic terminals labeled by the transport of neuronal tracers or substance P immunocytochemistry can form tubular clusters that surround the proximal dendrites of relay cells. In a rostral and lateral subdivision of the lateral LP nucleus (LPl-2), tectothalamic terminals form more typical glomerular arrangements. When compared with existing physiological data, these results suggest that a unique integration of tectal and cortical inputs may contribute to the response properties of LP neurons.
The mammalian pulvinar nucleus (PUL) establishes heavy interconnections with the parietal lobe, but the precise nature of these connections is only partially understood. To examine the distribution of corticopulvinar cells in the cat, we injected the PUL with retrograde tracers. Corticopulvinar cells were located in layers V and VI of a wide variety of cortical areas, with a major concentration of cells in area 7. To examine the morphology and distribution of corticopulvinar terminals, we injected cortical areas 5 or 7 with anterograde tracers. The majority of corticopulvinar axons were thin fibers (type I) with numerous diffuse small boutons. Thicker (type II) axons with fewer, larger boutons were also present. Boutons of type II axons formed clusters within restricted regions of the PUL. We examined corticopulvinar terminals labeled from area 7 at the ultrastructural level in tissue stained for γ-aminobutyric acid (GABA). By correlating the size of the presynaptic and postsynaptic profiles, we were able to quantitatively divide the labeled terminals into two categories: small and large (RS and RL, respectively). The RS terminals predominantly innervated small-caliber non-GABAergic (thalamocortical cell) dendrites, whereas the RL terminals established complex synaptic arrangements with dendrites of both GABAergic interneurons and non-GABAergic cells. Interpretation of these results using Sherman and Guillery's recent theories of thalamic organization (Sherman and Guillery [1998] Proc Natl Acad Sci U S A 95:7121-7126) suggests that area 7 may both drive and modulate PUL activity. Indexing termscortex; thalamus; visual system; sensorimotor; ultrastructure The feline pulvinar nucleus (PUL) receives input from a wide array of cortical areas (Raczkowski and Rosenquist, 1983) as well as the pretectum (PT;Berman, 1977;Berson and Graybiel, 1978;Schmidt et al., 2001;Baldauf et al., 2005). However, the contribution of these inputs to the response properties of PUL neurons is unknown. The cells of the PUL have large visual receptive fields that often lack clear boundaries; they respond more robustly to diffuse illumination than to small visual cues (Godfraind et al., 1972;Mason, 1981 by saccadic eye movements. Sudkamp and Schmidt (2000) identified three general classes of neurons in the feline PUL: "S" neurons are active during saccadic eye movements, "V" neurons are responsive to visual stimuli and unresponsive to eye movements, and "SV" neurons respond to both stationary ON and OFF stimuli and to sudden stimulus shifts.These response properties are similar in many respects to those of neurons in the parietal cortex, an area that establishes extensive reciprocal connections with the PUL (de V Clüver and Campos-Ortega, 1969;Heath and Jones, 1971;Robertson and Cunningham, 1981;Niimi et al., 1983;Raczkowski and Rosenquist, 1983;Avendaño et al., 1985;Olson and Lawler, 1987). In the cat, these cortical areas are primarily located within the middle suprasylvian gyrus (MSg) or cytoarchitectonically in areas 5 and 7 (Gu...
The distribution of the neuronal form of the nitric oxide-synthesizing enzyme, brain nitric oxide synthase (BNOS), was examined in the cat thalamus by using immunocytochemical techniques. BNOS was found in both cells and fibers throughout the visual thalamus. BNOS-stained cells were found consistently in the C laminae of the lateral geniculate nucleus (LGN), the pulvinar nucleus, and the lateral posterior nucleus (LP). In the A laminae of the LGN, variable numbers of BNOS-stained cells also could be detected. BNOS-stained cells were identified as a subset of interneurons because they all stained for glutamic acid decarboxylase (GAD), but not all GAD-stained cells contained BNOS. The average soma area of BNOS-stained cells was slightly greater than the average soma area of GAD-stained cells. BNOS-stained cells display a distinctive dendritic morphology, which is consistent with previous descriptions of class V neurons (Updyke [1979] J. Comp. Neurol. 186:603-619); they have widespread but fairly sparse arbors of thin, somewhat beaded dendrites. BNOS-stained cells participate in a distinct synaptic circuitry. Although many GAD-stained profiles are filled with vesicles and participate in complex synaptic arrangements, known as glomeruli, BNOS-stained dendrites contain small clusters of vesicles and form dendrodendritic contacts in the extraglomerular neuropil. Thus, there appear to be at least two types of gamma-aminobutyric acidergic interneurons in the visual thalamus of the cat. Interneurons that do not contain BNOS (class III morphology) may exert their effects primarily within synaptic glomeruli (Hamos et al. [1985] Nature 317:618-621), whereas interneurons that contain BNOS (class V morphology) contribute primarily to the extraglomerular neuropil.
One of the largest influences on dorsal lateral geniculate nucleus (dLGN) activity comes from interneurons, which use the neurotransmitter gamma-aminobutyric acid (GABA). It is well established that X retinogeniculate terminals contact interneurons and thalamocortical cells in complex synaptic arrangements known as glomeruli. However, there is little anatomical evidence for the involvement of dLGN interneurons in the Y pathway. To determine whether Y retinogeniculate axons contact interneurons, we injected the superior colliculus (SC) with biotinylated dextran amine (BDA) to backfill retinal axons, which also project to the SC. Within the A lamina of the dLGN, this BDA labeling allowed us to distinguish Y retinogeniculate axons from X retinogeniculate axons, which do not project to the SC. In BDA-labeled tissue prepared for electron microscopic analysis, we subsequently used postembedding immunocytochemical staining for GABA to distinguish interneurons from thalamocortical cells. We found that the majority of profiles postsynaptic to Y retinal axons were GABA-negative dendrites of thalamocortical cells (117/200 or 58.5%). The remainder (83/200 or 41.5%) were GABA-positive dendrites, many of which contained vesicles (59/200 or 29.5%). Thus, Y retinogeniculate axons do contact interneurons. However, these contacts differed from X retinogeniculate axons, in that triadic arrangements were rare. This indicates that the X and Y pathways participate in unique circuitries but that interneurons are involved in the modulation of both pathways.
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