Excitatory Projections of Wide Field Collicular Neurons to the Nucleus of the Optic Tract in the Rat.

The superficial layers of the mammalian superior colliculus (SC) contain neurons that are generally responsive to visual stimuli but can differ considerably in morphology and response properties. To elucidate the structure and function of these neurons, we combined extracellular recording and juxtacellular labeling, detailed anatomical reconstruction, and ultrastructural analysis of the synaptic contacts of labeled neurons, using transmission electron microscopy. Our labeled neurons project to different brainstem nuclei. Of particular importance are neurons that fit the morphological criteria of the wide field (WF) neurons and whose dendrites are horizontally oriented. They display a rather characteristic axonal projection pattern to the nucleus of optic tract (NOT); thus, we call them superior collicular WF projecting to the NOT (SCWFNOT) neurons. We corroborated the morphological characterization of this neuronal type as a distinct neuronal class with the help of unsupervised hierarchical cluster analysis. Our ultrastructural data demonstrate that SCWFNOT neurons establish excitatory connections with their targets in the NOT. Although, in rodents, the literature about the WF neurons has focused on their extensive projection to the lateral posterior nucleus of the thalamus, as a conduit for information to reach the visual association areas of the cortex, our data suggest that this subclass of WF neurons may participate in the optokinetic nystagmus.

Context dependence of head bobs in gerbils and potential neural contributions.

Headbobs are up-down movements of the cranium associated with the use of motion parallax for depth perception. Mongolian gerbils (aka jirds; Meriones unguiculatus) often execute a series of headbobs prior to jumping between surfaces. Gerbils were tested in a jumping stand task and headbobs videotaped under three light levels approximating low daylight, dawn/dusk, and moonlight across a range of distances to target. Headbobs per trial increased linearly with increasing distance to the target platform, whereas headbob frequency (rate of headbobbing pre-jump on the start platform) increased with gap distance up to an intermediate level and then decreased. Overall, gerbils made the most headbobs per trial under the darkest conditions, whereas their headbobbing rate was highest for medium illumination, especially for medium-long gap distances. There was a positive correlation between headbob frequency and volume of the superior colliculus (SC), but no relationship between headbobs and relative size of the temporo-posterior (TP) visual cortex. The results suggest that gerbils employ a specific visuomotor strategy for depth perception differentially under different conditions. We suggest that the deployment of headbobs under specific conditions may be part of an SC-driven vigilant state, of which more rapid sampling of the visual environment using headbobs for depth estimation is one component. Moreover, the findings highlight the importance of considering ecological factors in designing studies of visual behavior and its underpinnings in rodents.

Developmental distribution of primary cilia in the retinofugal visual pathway.

The mammalian visual system is composed of circuitry connecting sensory input from the retina to the processing core of the visual cortex. The two main retinorecipient brain targets, the superior colliculus (SC) and dorsal lateral geniculate nucleus (dLGN), bridge retinal input and visual output. The primary cilium is a conserved organelle increasingly viewed as a critical sensor for the regulation of developmental and homeostatic pathways in most mammalian cell types. Moreover, cilia have been described as crucial for neurogenesis, neuronal maturation, and survival in the cortex and retina. However, cilia in the visual relay center remain to be fully described. In this study, we characterized the ciliation profile of the SC and dLGN and found that the overall number of ciliated cells declined during development. Interestingly, shorter ciliated cells in both regions were identified as neurons, whose numbers remained stable over time, suggesting that cilia retention is a critical feature for optimal neuronal function in SC and dLGN. Our study suggests that primary cilia are important for neuronal maturation and function in cells of the SC and dLGN.

Anatomy and Connectivity of the Torus Longitudinalis of the Adult Zebrafish.

This study describes the cytoarchitecture of the torus longitudinalis (TL) in adult zebrafish by using light and electron microscopy, as well as its main connections as revealed by DiI tract tracing. In addition, by using high resolution confocal imaging followed by digital tracing, we describe the morphology of tectal pyramidal cells (type I cells) that are GFP positive in the transgenic line Tg(1.4dlx5a-dlx6a:GFP)ot1. The TL consists of numerous small and medium-sized neurons located in a longitudinal eminence attached to the medial optic tectum. A small proportion of these neurons are GABAergic. The neuropil shows three types of synaptic terminals and numerous dendrites. Tracing experiments revealed that the main efference of the TL is formed of parallel-like fibers that course within the marginal layer of the optic tectum. A toral projection to the thalamic nucleus rostrolateralis is also observed. Afferents to the TL come from visual and cerebellum-related nuclei in the pretectum, namely the central, intercalated and the paracommissural pretectal nuclei, as well as from the subvalvular nucleus in the isthmus. Additional afferents to the TL may come from the cerebellum but their origins could not be confirmed. The tectal afferent projection to the TL originates from cells similar to the type X cells described in other cyprinids. Tectal pyramidal neurons show round or piriform cell bodies, with spiny apical dendritic trees in the marginal layer. This anatomical study provides a basis for future functional and developmental studies focused on this cerebellum-like circuit in zebrafish.

Regional differences in the ependyma of the optic tectal ventricle of adult zebrafish with structures referring to brain hydrodynamics.

Classical electron microscopic morphological studies provide detailed ultrastructural information, which may lend insights into cellular functions. As a follow-up to our morphological investigation of the adult zebrafish (Danio rerio) optic tectum, in this study, we have analyzed the ependymal structures lining the surfaces of the tectal ventricle: the torus, tegmental surface of the valvula cerebelli and the periventricular gray zone of the optic tectal cortex. We used toluidine blue stained plastic (semithin) sections for light microscopy and scanning electron microscopy. Our morphological findings of gated entrances and/or egresses indicate that, at least in the adult zebrafish brain, there may be a bidirectional direct flow communication between the ventricular cerebrospinal fluid and the parenchymal interstitial fluid.

Retinorecipient areas in the diurnal murine rodent Arvicanthis niloticus: a disproportionally large superior colliculus.

The Nile grass rat (Arvicanthis niloticus) has a high proportion of cone photoreceptors (∼30-40%) compared with that in the common laboratory mouse and rat (∼1-3%) and may prove a preferable murine model with which to study cone-driven information processing in retina and primary visual centers. However, other than regions involved in circadian control, little is known about the retinorecipient structures in this rodent. We undertook a detailed analysis of the retinal projections as revealed after intravitreal injection of the anterograde tracer cholera toxin subunit B. Retinal efferents were evaluated in 45 subcortical structures. Contralateral projections were always dominant. Major contralateral inputs consisted of the suprachiasmatic nucleus, dorsolateral geniculate nucleus (dLGN), intergeniculate leaflet, ventral geniculate nucleus (magnocellular part), lateroposterior thalamic nucleus, all six pretectal nuclei, superficial layers of the superior colliculus (SC), and the main nuclei of the accessory optic system. Terminals from the contralateral eye were also localized in an unnamed field rostromedial to the dLGN as well as in the subgeniculate thalamic nucleus. Ipsilateral inputs were found mainly in the suprachiasmatic nucleus, dLGN, intergeniculate leaflet, internal sector of the magnocellular part of the ventral geniculate nucleus, olivary pretectal nucleus, and SC optic layer. Retinal afferents were not detected in the basal forebrain or the dorsal raphe nucleus. Morphometric measurements revealed that the superficial layers of the SC are disproportionately enlarged relative to other retinorecipient regions and brain size compared with rats and mice. We suggest that this reflects the selective projection of cone-driven retinal ganglion cells to the SC.

Control of excitatory CNS synaptogenesis by astrocyte-secreted proteins Hevin and SPARC.

Astrocytes regulate synaptic connectivity in the CNS through secreted signals. Here we identified two astrocyte-secreted proteins, hevin and SPARC, as regulators of excitatory synaptogenesis in vitro and in vivo. Hevin induces the formation of synapses between cultured rat retinal ganglion cells. SPARC is not synaptogenic, but specifically antagonizes synaptogenic function of hevin. Hevin and SPARC are expressed by astrocytes in the superior colliculus, the synaptic target of retinal ganglion cells, concurrent with the excitatory synaptogenesis. Hevin-null mice had fewer excitatory synapses; conversely, SPARC-null mice had increased synaptic connections in the superior colliculus. Furthermore, we found that hevin is required for the structural maturation of the retinocollicular synapses. These results identify hevin as a positive and SPARC as a negative regulator of synapse formation and signify that, through regulation of relative levels of hevin and SPARC, astrocytes might control the formation, maturation, and plasticity of synapses in vivo.

Topography of covert visual attention in human superior colliculus.

Experiments were performed to examine the topography of covert visual attention signals in human superior colliculus (SC), both across its surface and in its depth. We measured the retinotopic organization of SC to direct visual stimulation using a 90° wedge of moving dots that slowly rotated around fixation. Subjects (n = 5) were cued to perform a difficult speed-discrimination task in the rotating region. To measure the retinotopy of covert attention, we used a full-field array of similarly moving dots. Subjects were cued to perform the same speed-discrimination task within a 90° wedge-shaped region, and only the cue rotated around fixation. High-resolution functional magnetic resonance imaging (fMRI, 1.2 mm voxels) data were acquired throughout SC. These data were then aligned to a high-resolution T1-weighted reference volume. The SC was segmented in this volume so that the surface of the SC could be computationally modeled and to permit calculation of a depth map for laminar analysis. Retinotopic maps were obtained for both direct visual stimulation and covert attention. These maps showed a similar spatial distribution to visual stimulation maps observed in rhesus macaque and were in registration with each other. Within the depth of SC, both visual attention and stimulation produced activity primarily in the superficial and intermediate layers, but stimulation activity extended significantly more deeply than attention.

Visual deprivation increases accumulation of dense core vesicles in developing optic tectal synapses in Xenopus laevis.

Despite considerable progress in understanding the molecular components of synapses in the central nervous system, the ultrastructural rearrangements underlying synaptic development remain unclear. We used serial section transmission electron microscopy and three-dimensional reconstructions of the optic tectal neuropil of Xenopus laevis tadpoles to detect and quantify changes in synaptic ultrastructure over a 1-week period from stages 39 and 47, during which time the visual system of Xenopus tadpoles becomes functional. Synapse density, presynaptic maturation index, and number of synapses per axon bouton increase, whereas the number of DCVs per bouton decreases, between stages 39 and 47. The width of the synaptic cleft decreased and the diameter of postsynaptic profiles increased between stages 39 and 47 and then remained relatively unchanged after stage 47. We found no significant difference in synapse maturation between GABAergic and non-GABAergic synapses. To test the effect of visual experience on synaptogenesis, animals were deprived of visual experience for 3 days from stage 42 to 47. Visual deprivation decreased synapse maturation and the number of connections per bouton. Furthermore, visual deprivation increased the number of DCVs per bouton by more than twofold. The visual-deprivation-induced decrease in synaptic connections is specific to asymmetric non-GABAergic synapses; however, both symmetric GABAergic and asymmetric synapses show comparable increases in the number DCVs with visual deprivation. In both the control and the visually deprived animals, the number of DCVs per bouton is highly variable and does not correlate with either synapse maturation or the number of connected partners per bouton. These data suggest that synaptogenesis and DCV accumulation are regulated by visual experience and further suggest a complex spatial and temporal relation between DCV accumulation and synapse formation.

Laminar restriction of retinal ganglion cell dendrites and axons: subtype-specific developmental patterns revealed with transgenic markers.

Retinal ganglion cells (RGCs), which transfer information from the eye to the brain, are heterogeneous in structure and function, but developmental studies have generally treated them as a single group. Here, we investigate the development of RGC axonal and dendritic arbors using four mouse transgenic lines in which nonoverlapping subsets of RGCs are indelibly labeled with a fluorescent protein. Each subset has a distinct functional signature, size, and morphology. Dendrites of each subset are restricted to specific sublaminae within the inner plexiform layer in adulthood, but acquire their restriction in different ways: one subset has lamina-restricted dendrites from an early postnatal stage, a second remodels an initially diffuse pattern, and two others develop stepwise. Axons of each subset arborize in discrete laminar zones within the lateral geniculate nucleus or superior colliculus, demonstrating previously unrecognized subdivisions of retinorecipient layers. As is the case for dendrites, lamina-restricted axonal projections of RGC subsets develop in different ways. For example, while axons of two RGC subsets arborize in definite zones of the superior colliculus from an early postnatal stage, axons of another subset initially occupy a deep layer, then translocate to a narrow subpial zone. Together, these results show that RGC subsets use a variety of strategies to construct lamina-restricted dendritic and axonal arbors. Taking account of these subtype-specific features will facilitate identification of the molecules and cells that regulate arbor formation.

The macaque midbrain reticular formation sends side-specific feedback to the superior colliculus.

The central mesencephalic reticular formation (cMRF) likely plays a role in gaze control, as cMRF neurons receive tectal input and provide a bilateral projection back to the superior colliculus (SC). We examined the important question of whether this feedback is excitatory or inhibitory. Biotinylated dextran amine (BDA) was injected into the cMRF of M. fascicularis monkeys to anterogradely label reticulotectal terminals and retrogradely label tectoreticular neurons. BDA labeled profiles in the ipsi- and contralateral intermediate gray layer (SGI) were examined electron microscopically. Postembedding GABA immunochemistry was used to identify putative inhibitory profiles. Nearly all (94.7%) of the ipsilateral BDA labeled terminals were GABA positive, but profiles postsynaptic to these labeled terminals were exclusively GABA negative. In addition, BDA labeled terminals were observed to contact BDA labeled dendrites, indicating the presence of a monosynaptic feedback loop connecting the cMRF and ipsilateral SC. In contrast, within the contralateral SGI, half of the BDA labeled terminals were GABA positive, while more than a third were GABA negative. All the postsynaptic profiles were GABA negative. These results indicate the cMRF provides inhibitory feedback to the ipsilateral side of the SC, but it has more complex effects on the contralateral side. The ipsilateral projection may help tune the "winner-take-all" mechanism that produces a unified saccade signal, while the contralateral projections may contribute to the coordination of activity between the two colliculi.

[Evolutionary significance of reciprocal connections in the turtle tectofugal visual system].

In two turtle species--Emys orbicularis and Testudo horsfieldi--by the method of anterograde and retrograde traicing method at the light and electron microscopy level, the existence is proven of direct descending projections from the thalamic nucleus of the tectofugal visual system n. rotunds (Rot) to the optic tectum. After injection of tracers into Rot alone and into Rot with involvement of the tectothalamic tract (Trtth), occasional labeled fibers with varicosities and terminals are revealed predominantly in the deep sublayers of SGFS of the rostral optic tectum, while in the lower amount in other tectal layers. After the tracer injections into the optic tectum, a few retrogradely labeled neurons were found mainly in the Rot ventral parts and within Trtth. Their localization coincides with that of GABA-immunoreactive cells. Electron microscopy showed the existence of many retrogradely labeled dendrites throughout the whole Rot; a few labeled cell bodies were also present there, some of them being also GABA-immunoreactive. These results allow us to conclude about the existence of reciprocal connections between the optic tectum and Rot in turtles, these connections being able to affect processing of visual information in tectum. We suggest that reciprocity of tectothalamic connections might be the ancestral feature of the vertebrate brain; in the course of amniote evolution the functional significance of this feature can be decreased and even lost in parallel with a rise of the role of direct corticotectal projections.

Short-latency visual input to the subthalamic nucleus is provided by the midbrain superior colliculus.

The subthalamic nucleus (STN) is one of the principal input nuclei of the basal ganglia. Using electrophysiological techniques in anesthetized rats, we show that the STN becomes responsive to visual stimuli at short latencies when local disinhibitory injections are made into the midbrain superior colliculus (SC), an important subcortical visual structure. Significantly, only injections into the lateral, but not medial, deep layers of the SC were effective. Corresponding disinhibition of primary visual cortex also was ineffective. Complementary anatomical analyses revealed a strong, regionally specific projection from the deep layers of the lateral SC to neurons in rostral and dorsal sectors of the STN. Given the retinocentric organization of the SC, these results suggest that lower-field stimuli represented in the lateral colliculus have a direct means of communicating with the basal ganglia via the STN that is not afforded to visual events occurring in the upper visual field.

Contrasting the functional properties of GABAergic axon terminals with single and multiple synapses in the thalamus.

Diverse sources of GABAergic inhibition are a major feature of cortical networks, but distinct inhibitory input systems have not been systematically characterized in the thalamus. Here, we contrasted the properties of two independent GABAergic pathways in the posterior thalamic nucleus of rat, one input from the reticular thalamic nucleus (nRT), and one "extrareticular" input from the anterior pretectal nucleus (APT). The vast majority of nRT-thalamic terminals formed single synapses per postsynaptic target and innervated thin distal dendrites of relay cells. In contrast, single APT-thalamic terminals formed synaptic contacts exclusively via multiple, closely spaced synapses on thick relay cell dendrites. Quantal analysis demonstrated that the two inputs displayed comparable quantal amplitudes, release probabilities, and multiple release sites. The morphological and physiological data together indicated multiple, single-site contacts for nRT and multisite contacts for APT axons. The contrasting synaptic arrangements of the two pathways were paralleled by different short-term plasticities. The multisite APT-thalamic pathway showed larger charge transfer during 50-100 Hz stimulation compared with the nRT pathway and a greater persistent inhibition accruing during stimulation trains. Our results demonstrate that the two inhibitory systems are morpho-functionally distinct and suggest and that multisite GABAergic terminals are tailored for maintained synaptic inhibition even at high presynaptic firing rates. These data explain the efficacy of extrareticular inhibition in timing relay cell activity in sensory and motor thalamic nuclei. Finally, based on the classic nomenclature and the difference between reticular and extrareticular terminals, we define a novel, multisite GABAergic terminal type (F3) in the thalamus.

Orexin-A immunoreactive cells and fibers in the central nervous system of the axolotl brain and their association with tyrosine hydroxylase and serotonin immunoreactive somata.

Orexin-A-like immunoreactivity in the axolotl brain was investigated by immunohistochemistry. Immunoreactive somata formed a single group in the hypothalamus, but were distributed beyond several nuclei, namely, the ventral aspect of the nucleus preopticus posterior, dorsal aspect of the nucleus suprachiasmaticus and anterior aspect of the pars ventralis hypothalami. Immunoreactive fibers were distributed throughout the brain from the olfactory bulb to the spinal cord except the cerebellum. The densest immunoreactive fibers were seen in the medial forebrain bundle and caudal lateral forebrain bundle. The largest number of immunoreactive puncta were seen in the mesencephalic tectum in addition to the hypothalamus. Immunoelectron microscopic analysis revealed the presence of synaptoid connections of immunoreactive fibers on neuronal somata in the tectum. The function of the mesencephalic system in the urodele seems to be sensory integration, suggesting that the orexin-A nervous system is associated with the modulation of sensory inputs. Orexin-A immunoreactive puncta were also observed on catecholaminergic and serotonergic somata. In view of the restricted somatic distribution in the hypothalamus, wide distribution of fibers throughout the central nervous system (CNS), and intimate association with monoaminergic somata, the orexin nervous system in the axolotl CNS is similar to those of other vertebrates, suggesting that this system is essential for brain functions throughout vertebrates.

Tectorotundal connections in turtles: an electron microscopic tracing and GABA-immunocytochemical study.

The nucleus rotundus of the turtles Emys orbicularis and Testudo horsfieldi was analysed by axonal tracing methods and post-embedding GABA immunocytochemistry. After injections of horseradish peroxidase or biotinylated dextran amine into the optic tectum, electron microscopic observations showed that the vast majority of ipsilateral tectorotundal axon terminals were small in size, had smooth contours and contained small, round, densely packed synaptic vesicles. These terminals were GABA-immunonegative, often gathered in clusters, and established asymmetrical synaptic contacts with either small- or medium-sized GABA-negative dendritic profiles and with GABA-immunoreactive (GABA-ir) dendrites, which did not contain synaptic vesicles. Occasional GABA-ir-labelled axon terminals were observed; these may arise from the rare GABAergic neurons in the central tectal layer, or from neurons in the ventral pretectal nucleus, which projects both to the optic tectum and nucleus rotundus. In addition to tracer-labelled axon terminals, we observed both GABA-negative and GABA-ir cell bodies and dendrites also labelled by the tracer. No GABA-ir presynaptic dendritic profiles containing synaptic vesicles were observed. The existence in reptiles of reciprocal connections between the nucleus rotundus and the optic tectum as a phylogenetically ancient feedback system is discussed.

Anterograde axonal transport of chicken cellular prion protein (PrPc) in vivo requires its N-terminal part.

The cellular isoform of prion protein (PrP(c)) can exist in membrane-bound and secreted forms. Both forms of PrP(c) can be transported by retinal ganglion cell (RGC) axons along the optic nerve in the anterograde direction. In this study we determined which part of chicken PrP(c) is required for its anterograde axonal transport within the optic nerve of embryonic chicken. We intraocularly injected radio-iodinated fragments of recombinant chicken PrP(c) and then examined their anterograde axonal transport from retina into optic tectum. Using gamma-counting and different autoradiographic techniques we quantified anterograde axonal transport of the N-terminal part of chicken PrP(c) (amino acid residues 1-116) in this model system. The transport of the N-terminal part has similar properties as the anterograde transport of full-length chicken PrP(c) (Butowt et al., 2006) described previously (e.g., has similar efficiency, is microtubule-dependent, and is saturable). Moreover, the pattern of ultrastructural distribution of the N-terminal fragment within RGCs is similar to the distribution of full-length PrP(c). The C-terminal fragment of chicken PrP(c) (residues 118-246) and different PrP-derived peptides were not transported. Moreover, PrP(c)-derived peptides were sorted into different endocytotic pathways in neurons, indicating that they cannot substitute for full-length PrP(c) to study its internalization and trafficking. These data indicate that the N-terminal half of chicken PrP(c) contains the necessary information to drive the internalization and subsequent sorting of extracellular PrP(c) in RGCs soma into the anterograde axonal transport pathway.

Cell-autonomous TrkB signaling in presynaptic retinal ganglion cells mediates axon arbor growth and synapse maturation during the establishment of retinotectal synaptic connectivity.

BDNF contributes to the activity-dependent establishment and refinement of visual connectivity. In Xenopus, BDNF applications in the optic tectum influence retinal ganglion cell (RGC) axon branching and promote synapse formation and stabilization. The expression patterns of BDNF and TrkB suggest that BDNF specifically regulates the maturation of RGC axons at the target. It is possible, however, that BDNF modulates retinotectal synaptic connectivity by differentially influencing presynaptic RGC axons and postsynaptic tectal cells. Here, we combined single-cell expression of a dominant-negative TrkB-enhanced green fluorescent protein (GFP) fusion protein with confocal microscopy imaging in live Xenopus tadpoles to differentiate between presynaptic and postsynaptic actions of BDNF. Disruption of TrkB signaling in individual RGCs influenced the branching and synaptic maturation of presynaptic axon arbors. Specifically, GFP-TrkB.T1 overexpression increased the proportion of axons with immature, growth cone-like morphology, decreased axon branch stability, and increased axon arbor degeneration. In addition, GFP-TrkB.T1 overexpression reduced the number of red fluorescent protein-synaptobrevin-labeled presynaptic specializations per axon terminal. In contrast, overexpression of GFP-TrkB.T1 in tectal neurons did not alter synaptic number or the morphology or dynamic behavior of their dendritic arbors. Electron microscopy analysis revealed a significant decrease in the number of mature synaptic profiles and in the number of docked synaptic vesicles at retinotectal synapses made by RGC axons expressing GFP-TrkB.T1. Together, our results demonstrate that presynaptic TrkB signaling in RGCs is a key determinant in the establishment of visual connectivity and indicate that changes in tectal neuron synaptic connectivity are secondary to the BDNF-elicited enhanced stability and growth of presynaptic RGCs.

The superficial layers of the superior colliculus are cytoarchitectually and myeloarchitectually disorganized in the reelin-deficient mouse, reeler.

The causative gene for the reeler mouse is reelin which encodes Reelin protein, an extracellular molecule. In the present study, we have examined the cytoarchitecture, myeloarchitecture, and afferent/efferent systems of the superior colliculus (SC) of the reeler mouse. In the reeler, the laminar structures of the superficial three layers of the SC were disorganized and intermingled into a single layer, i.e., the superficial fused layer (SuF), as previously reported in the reelin-deficient SRK rat (Sakakibara et al., Develop. Brain Res. 141:1-13). Next, we have investigated the course and terminals of visual corticotectal and retinotectal projections with an injection of biocytin into the visual cortex or an injection of cholera toxin subunit B into the retina, respectively. In the reeler, anterogradely labeled visual corticotectal and retinotectal fibers took an aberrant course within the SuF, resulting in abnormal myeloarchitecture of the superficial SC of the reeler. Retrograde labeling of tectospinal tract neurons could not show any differences between the normal and reeler mice, suggesting that the deep layers of the reeler SC are cytoarchitectually normal. In situ hybridization and immunohistochemical studies have shown that reelin mRNA and Reelin protein were both recognized in the normal SC. These results suggest that Reelin protein plays some roles in histogenesis of the superficial layers of the SC.

Comparison of the ultrastructure of cortical and retinal terminals in the rat superior colliculus.

We compared the ultrastructure and synaptic targets of terminals of cortical or retinal origin in the stratum griseum superficiale and stratum opticum of the rat superior colliculus. Following injections of biotinylated dextran amine into cortical area 17, corticotectal axons were labeled by anterograde transport. Corticotectal axons were of relatively small caliber with infrequent small varicosities. At the ultrastructural level, corticotectal terminals were observed to be small profiles (0.44 +/- 0.27 microm(2)) that contained densely packed round vesicles. In tissue stained for gamma amino butyric acid (GABA) using postembedding immunocytochemical techniques, corticotectal terminals were found to contact small (0.51 +/- 0.69 microm(2)) non-GABAergic dendrites and spines (93%) and a few small GABAergic dendrites (7%). In the same tissue, retinotectal terminals, identified by their distinctive pale mitochondria, were observed to be larger than corticotectal terminals (3.34 +/- 1.79 microm(2)). In comparison to corticotectal terminals, retinotectal terminals contacted larger (1.59 +/- 1.70 microm(2)) non-GABAergic dendrites and spines (73%) and a larger proportion of GABAergic profiles (27%) of relatively large size (2.17 +/- 1.49 microm(2)), most of which were vesicle-filled (71%). Our results suggest that cortical and retinal terminals target different dendritic compartments within the neuropil of the superficial layers of the superior colliculus.