Two pathways from the superior colliculus (SC) to the tree shrew pulvinar nucleus have been described, one in which the axons terminate in dense (or specific) patches and one in which the axon arbors are more diffusely organized J. . As predicted by Lyon et al. ([2003] J. Comp. Neurol. 467:593-606), we found that anterograde labeling of the diffuse tectopulvinar pathway terminated in the acetylcholinesterase (AChE)-rich dorsal pulvinar (Pd), whereas the specific pathway terminated in the AChE-poor central pulvinar (Pc). Injections of retrograde tracers in Pd labeled non-γ-aminobutyric acid (GABA)-ergic wide-field vertical cells located in the lower stratum griseum superficiale and stratum opticum of the medial SC, whereas injections in Pc labeled similar cells in more lateral regions. At the ultrastructural level, we found that tectopulvinar terminals in both Pd and Pc contact primarily non-GABAergic dendrites. When present, however, synaptic contacts on GABAergic profiles were observed more frequently in Pc (31% of all contacts) compared with Pd (16%). Terminals stained for the type 2 vesicular glutamate transporter, a potential marker of tectopulvinar terminals, also contacted more GABAergic profiles in Pc (19%) compared with Pd (4%). These results provide strong evidence for the division of the tree shrew pulvinar into two distinct tectorecipient zones. The potential functions of these pathways are discussed. J. Comp. Neurol. 510:24 -46, 2008. Indexing termssynapse; ultrastructure; GABA; pulvinar nucleus; superior colliculus; vesicular glutamate transporter; Tupaia belangeri Parallel visual pathways from the retina to the cortex, relayed via the dorsal lateral geniculate nucleus (dLGN), or the superior colliculus (SC) and pulvinar nucleus, likely serve distinct functions in the coding of form, movement, and spatial location signals. In the dLGN, further segregations of anatomically and physiologically distinct visual pathways have been identified and extensively characterized (Sherman, 1985). Likewise, studies in a variety of species have provided evidence for the existence of multiple pathways from the SC to the thalamus (May, 2006), although these pathways are largely uncharacterized, and their functions are unclear. The tree shrew, with its expanded tectopulvinar system, is good choice for studies of how pathways from the SC influence cortical activity via their projections to the pulvinar nucleus.In 1988, Luppino et al. labeled tectothalamic terminals in the tree shrew by placing small injections of axonal tracers in the SC and discovered that the pulvinar nucleus receives input *Correspondence to: Martha E. Bickford, Department of Anatomical Sciences and Neurobiology, University of Louisville, School of Medicine, 500 S. Preston St., Louisville, KY 40292. E-mail: martha.bickford@louisville.edu. We recently examined the synaptic organization of two tectorecipient zones of the cat thalamus (Kelly et al., 2003). We found that tectal terminals in the medial subdivision of the lateral posterior (LPm) ...
Visually guided movement is possible in the absence of conscious visual perception, a phenomenon referred to as “blindsight.” Similarly, fearful images can elicit emotional responses in the absence of their conscious perception. Both capabilities are thought to be mediated by pathways from the retina through the superior colliculus (SC) and pulvinar nucleus. To define potential pathways that underlie behavioral responses to unperceived visual stimuli, we examined the projections from the pulvinar nucleus to the striatum and amygdala in the tree shrew (Tupaia belangeri), a species considered to be a prototypical primate. The tree shrew brain has a large pulvinar nucleus that contains two SC-recipient subdivisions; the dorsal (Pd) and central (Pc) pulvinar both receive topographic (“specific”) projections from SC, and Pd receives an additional non-topographic (“diffuse”) projection from SC (Chomsung et al., 2008). Anterograde and retrograde tract tracing revealed that both Pd and Pc project to the caudate and putamen, and Pd, but not Pc, additionally projects to the lateral amygdala. Using immunocytochemical staining for substance P (SP) and parvalbumin (PV) to reveal the patch/matrix organization of tree shrew striatum, we found that SP-rich/PV-poor patches interlock with a PV-rich/SP-poor matrix. Confocal microscopy revealed that tracer-labeled pulvino-striatal terminals preferentially innervate the matrix. Electron microscopy revealed that the postsynaptic targets of tracer-labeled pulvino-striatal and pulvino-amygdala terminals are spines, demonstrating that the pulvinar nucleus projects to the spiny output cells of the striatum matrix and the lateral amygdala, potentially relaying: (1) topographic visual information from SC to striatum to aid in guiding precise movements, and (2) non-topographic visual information from SC to the amygdala alerting the animal to potentially dangerous visual images.
1. This paper represents a continuation of our effort to examine the relationship between the physiology of distinct classes of primate lateral geniculate nucleus (LGN) cells and spatial vision. The present study focuses on modeling the contrast-sensitivity functions (CSFs) of separate LGN cell classes, examining differences in the CSFs of different classes of LGN cells and comparing the results with behaviorally defined CSFs. 2. CSFs to drifting sinusoidal gratings were obtained from single LGN relay cells in the nocturnal primate, Galago crassicaudatus. The CSFs of 14 X-like, 27 Y-like, and 6 W-like cells with standard center-surround organization were well fit by a difference of Gaussians (DOG) model with small residual errors (mean error per data point +/- SEM = 0.008 +/- 0.002). The larger residual errors shown by a few of the Y-like cells were not due to nonlinearity of spatial summation. 3. The CSFs of eight cells that appeared to have nonstandard center-surround organization (primarily, a silent, suppressive surround) were also well fit by the DOG model. 4. The DOG curves that best fitted the data differed considerably between the three groups. As a group, X-like cells had a small center mechanism (Rc = 0.19 degrees) with high sensitivity (Kc = 76.53) and a small, sensitive surround (Rs = 0.71 degrees; Ks = 5.50). These parameters produced CSFs with high cutoff frequencies (Vcutoff = 2.5 c/deg) and low peak sensitivities (CSpk = 6.1) that occurred at 0.8 c/deg. 5. Y-like cells had a large center mechanism (Rc = 0.46 degrees) with low sensitivity (Kc = 21.16) and a large, insensitive surround (Rs = 2.38 degrees; Ks = 0.81). These parameters produced CSFs with lower cutoff frequencies (Vcutoff = 1.2 c/deg) and higher peak sensitivities (CSpk = 12.5) that occurred at 0.2 c/deg. 6. The few W-like cells that responded to gratings well enough to determine a CSF were quite variable. As a group they had a large center mechanism (Rc = 0.38 degrees) with intermediate sensitivity (Kc = 34.55) and a surround with intermediate size and sensitivity (Rs = 1.59 degrees; Ks = 1.59). These produced CSFs with intermediate cutoffs (Vcutoff = 1.6 c/deg) and low peak sensitivities (CSpk = 5.0) occurring at 0.4 c/deg.(ABSTRACT TRUNCATED AT 400 WORDS)
We examined the synaptic organization of reciprocal connections between the temporal cortex and the dorsal (Pd) and central (Pc) subdivisions of the tree shrew pulvinar nucleus, regions innervated by the medial and lateral superior colliculus, respectively. Both Pd and Pc subdivisions project topographically to 2 separate regions of the temporal cortex; small injections of anterograde tracers placed in either Pd or Pc labeled 2 foci of terminals in the temporal cortex. Pulvinocortical pathways innervated layers I-IV, with beaded axons oriented perpendicular to the cortical surface, where they synapsed with spines that did not contain gamma amino butyric acid (GABA), likely located on the apical dendrites of pyramidal cells. Projections from the temporal cortex to the Pd and Pc originate from layer VI cells, and form small terminals that contact small caliber non-GABAergic dendrites. These results suggest that cortical terminals are located distal to tectopulvinar terminals on the dendritic arbors of Pd and Pc projection cells, which subsequently contact pyramidal cells in the temporal cortex. This circuitry could provide a mechanism for the pulvinar nucleus to activate subcortical visuomotor circuits and modulate the activity of other visual cortical areas. The potential relation to primate tecto-pulvino-cortical pathways is discussed.
Relay neurons in dorsal thalamic nuclei can fire high frequency bursts of action potentials that ride the crest of voltage-dependent transient (T-type) calcium currents (low threshold spike; LTS). To explore potential nucleus-specific burst features, we compared the membrane properties of dorsal lateral geniculate nucleus (dLGN) and pulvinar nucleus relay neurons using in vitro whole cell recording in juvenile and adult tree shrew (Tupaia) tissue slices. We injected current ramps of variable slope into neurons that were sufficiently hyperpolarized to de-inactivate T-type calcium channels. In a small percentage of juvenile pulvinar and dLGN neurons, an LTS could not be evoked. In the remaining juvenile neurons, and in all adult dLGN neurons, a single LTS could be evoked by current ramps. However, in the adult pulvinar, current ramps evoked multiple LTSs in over 70% of recorded neurons. Using immunohistochemistry, western blot techniques, unbiased stereology, confocal and electron microscopy, we found that pulvinar neurons expressed more T-type calcium channels (Cav 3.2) and more small conductance potassium channels (SK2) than dLGN neurons and that the pulvinar nucleus contained a higher glia-to-neuron ratio than the dLGN. Hodgkin-Huxley type compartmental models revealed that the distinct firing modes could be replicated by manipulating T-type calcium and SK2 channel density, distribution, and kinetics. The intrinsic properties of pulvinar neurons that promote burst firing in the adult may be relevant to the treatment of conditions that involve the adult onset of aberrant thalamocortical interactions.
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