The morphology of corticofugal axons to the dorsal lateral geniculate nucleus in the cat

The structural features of corticogeniculate axons were studied in adult cats after labeling them with horseradish peroxidase (HRP). Injections of HRP into the optic radiations near the dorsal lateral geniculate nucleus result in Golgi-like filling of both geniculate relay neurons and corticogeniculate axons. In the present material at least two main types of axons could be defined. The most common type is called the type I axon because it so closely resembles the type I axons described by Guillery ('66, '67) in Golgi preparations. These fine axons have smooth surfaces and consistent fiber diameter. Most terminal swellings are at the ends of short collateral branches and these swellings form asymmetric synaptic contacts onto small and medium-sized dendrites. Type I axons typically innervate more than one lamina as well as interlaminar zones and they clearly arise from the cerebral cortex. The second type of axon is called the beaded axon because of its numerous swellings, en passant. These swellings frequently are larger than those on type I axons and they differ from previously described corticogeniculate axon terminals in their ultrastructural features. That is, their synaptic contacts appear symmetrical and they form axosomatic contacts. Because of these differences, the possibility that beaded axons are of subcortical origin, particularly from the perigeniculate nucleus, is discussed. When type I axons and geniculate relay neurons are filled in the same region of the nucleus it is possible to identify probable sites of synaptic contact by using the light microscope. Such analyses indicate that corticogeniculate axons synapse directly onto relay cells, primarily on peripheral dendritic branches. Further, it appears that single axons contact many geniculate neurons and that single neurons are contacted by many axons.     

Electrophysiological and morphological properties of neurons in the ventral lateral geniculate nucleus of the rabbit

The ventral lateral geniculate nucleus of the rabbit has been studied with light and electron microscopy, as well as single unit extracellular recordings. There are two basic types of neurons in the nucleus, and three synaptic types, designated RS, RL and F. There are also synaptic glomeruli, in which RL axons are always involved. The arrangement of these synaptic types and their percentage of occurrence closely resembles that in dorsal thalamic nuclei. Retinal projections are found in both external and internal sectors of the nucleus, shown by both degeneration and autoradiographic techniques. Retinal projections to the internal sector are sparser and degenerate more rapidly than those to the external sector. Electrophysiological studies revealed some neurons in both external and internal sectors with receptive fields resembling those in the dorsal lateral geniculate nucleus. There were, however, a significant number of neurons with an indefinite visual response or no visual response at all. The possible role of the ventral lateral geniculate nucleus in visual function is discussed.     

The contribution of the dorsal lateral geniculate nucleus to the total pattern of thalamic terminations in striate cortex of the Virginia opossum

These studies were carried out to show the manner of projection of the dorsal lateral geniculate nucleus and other thalamic nuclei to striate cortex in the Virginia opossum. In order to demonstrate these projections, lesions were made in the dorsal lateral geniculate nucleus, in the ventral lateral geniculate nucleus, in most of the thalamus on one side except for the dorsal lateral geniculate nucleus, and in the entire unilateral thalamus. Following various survival times, usually seven days, the brains were appropriately prepared and stained with procedure I of the Fink-Heimer technique. Dorsal lateral geniculate neurons project in a topographical manner only to certain layers of striate cortex. These projections from the lateral geniculate are compared with the same system in other mammals, and it is concluded that it is similar in all mammals studied, except for the cat. In the cat the lateral geniculate projects beyond the border of striate cortex, but even in the cat the layers of termination within striate cortex are apparently similar. The ventral lateral geniculate nucleus does not project to visual cortex. Dorsal thalamic nuclei other dian the lateral geniculate project to peristriate cortex and to layers VI and I of striate cortex. The finding that thalamic nuclei, other than the lateral geniculate nucleus, project to striate cortex has never been described as part of the visual pathways in other mammals. It is suggested that these additional projections arise mainly from the lateral nuclear group of the thalamus in the opossum, and must be considered in relation to any response characteristics and organization of striate cells determined from physiological studies. These multiple thalamic projections can provide the substrate for more than one representation or “map” of sensory information in striate cortex.     

Organization of cholinergic synapses in the cat's dorsal lateral geniculate and perigeniculate nuclei

In the preceding article, we showed that cholinergic fibers originating from the brainstem reticular formation provide a dense innervation of the lateral geniculate nucleus. In this report we describe the ultrastructure of these fibers and their relations with other elements in the neuropil of the lateral geniculate nucleus. Cholinergic fibers were labeled with an antibody to choline acetyltransferase (ChAT), the synthesizing enzyme for acetylcholine (ACh). In the A-laminae of the lateral geniculate nucleus, ChAT + profiles are small and contain tightly packed, mostly round vesicles. Some end in encapsulated synaptic zones where they form asymmetrical synaptic contacts with processes of both projection cells and interneurons. Others form synapses upon the shafts of dendrites. Of the four classical types of vesicle-containing profiles identified by Guillery (Z. Zellforsch. Mikrosk. 96:1-38, '69; Vision Res. [Suppl.] 3:211-227, '71), ChAT + profiles most closely resemble RSD profiles (Round vesicles, Small profile, Dark mitochondria). However, as a population, ChAT + profiles can be distinguished from the unlabeled population of RSD profiles because they are larger in size, contain more mitochondria, and make synapses with smaller postsynaptic membrane specializations. Each of these differences is statistically significant and together they indicate that ChAT + profiles are a distinct morphological type of synaptic profile. ChAT + profiles in the perigeniculate nucleus resemble those found in the lateral geniculate nucleus; they also make synapses with obvious postsynaptic thickenings.     

Synaptic connections between corticogeniculate axons and interneurons in the dorsal lateral geniculate nucleus of the cat

Anatomical evidence is provided for direct synaptic connections by axons from visual cortex with interneurons in lamina A of the cat's dorsal lateral geniculate nucleus. Corticogeniculate axon terminals were labeled selectively with 3H-proline and identified by means of electron microscopic autoradiography. Interneurons in the lateral geniculate nucleus were stained with antibodies that had been raised against gamma aminobutyric acid (GABA). We found that corticogeniculate terminals synapsed with dendrites stained positively for GABA about three times as often as with unstained dendrites. Of the corticogeniculate terminals that contacted GABA-positive dendrites, 97% made synaptic connections with dendritic shafts. Only 3% synapsed with F profiles, the vesicle-filled dendritic appendages characteristic of lateral geniculate interneurons. These results suggest that the corticogeniculate pathway in the cat is directed primarily at interneurons and is organized synaptically to influence the integrated output of these cells, rather than the local interactions in which their dendritic specializations participate.     

The organization of the pulvinar in the grey squirrel (Sciurus carolinensis). II. Synaptic organization and comparisons with the dorsal lateral geniculate nucleus

The purpose of these experiments was to compare the synaptic organization of the subdivisions of the pulvinar defined in the preceding paper (Robson and Hall, '77) with each other and with the organization present in the dorsal lateral geniculate nucleus. The electron microscope was used to analyze normal synaptic arrangements and degenerating axonal terminals resulting from lesions. The dorsal lateral geniculate nucleus in the grey squirrel contains synaptic clusters similar to those described previously for other species. These clusters are characterized by large optic tract terminals which form multiple contacts onto large dendritic processes and other processes containing flat or pleomorphic vesicles. The geniculate lamina adjacent to the optic tract receives projections from the superior colliculus as well are from the retina. The terminals of the superior colliculus axons are small and medium sized and lie outside of the synaptic clusters. The retinal terminals are in the clusters. In the pulvinar, the rostro-medial subdivision contains synaptic clusters which resemble those in the lateral geniculate nucleus. These clusters contain large axon terminals which make multiple contacts onto large dendrites. However, these terminals are not contributed by an ascending sensory pathway but by axons from strait cortex. The rostro-lateral and caudal subdivisions of the pulvinar also contain synaptic clusters, but these clusters consist of a segment of a large dendrite which is ensheathed by medium-sized terminals. Since only a few of these medium sized terminals in any one cluster degenerate after tectal lesions, and none degenerate after cortical lesions, it is suggested that the morphological arrangement of these clusters may permit the convergence of axons from several sources, some of which are unidentified, onto the same dendritic segment.     

Thalamo-cortical organization of the visual system in the cat

Thalamo-cortical relationships in the visual system of the cat were studied by the method of retrograde degeneration. Localized lesions limited to area 17 result in degeneration only in the dorsolateral geniculate body; cell changes are marked in 3 laminae (A, A₁, B), mild in nucleus interlaminaris centralis and minimal in nucleus interlaminaris medialis. Lesions limited to areas 18 and 19 are followed by marked degeneration in the medial interlaminar nucleus, mild in the other laminae; in addition, the lateral part of the posterior thalamic nucleus (ventral or inferior pulvinar) is also atrophied. Following large striate lesions which marginally involved areas 18 and 19, there is also mild, localozed degeneration in the anteroventral and reticular thalamic nuclei. Whin cortical lesions are limited to the convexity of the suprasylvian gyri, degeneration is present in the lateral aspect of laminae A, A₁, B and nucleus interlaminaris centralis and in the medial part of the posterior nucleus, in addition to lateral dorsal, lateral posterior and pulvinar nuclei. Lesions in the ectosylvian gyri result in slight but definite degeneration in the lateral part of lamina A of the dorsal lateral geniculate, but nothing in the posterior nucleus. The geniculate projections to areas 17, 18 and 19, to the suprasylvian and ectosylvian gyri all show a rostrocaudal organization. The geniculostriate projection is also topographically organized in a mediolateral manner. Thus, the geniculocortical projection in the cat is not striate specific but spreads over the occipito-temporal cortex at least as far as the acoustic areas of the ectosylvian gyri. In this species the dorsal lateral geniculate body is not a unitary structure but is a complex of nuclei, all of which receive retinal fibers, and the cortical projections of which overlap those of the posterior, lateral dorsal, lateral posterior, pulvinar, medial geniculate, reticular and anterior thalamic nuclei.     

Auditory pathways to the cortex in Tupaia glis

The auditory system of the tree shrew, Tupaia glis, was investigated by identifying axonal degeneration after lesions of the lateral lemniscus, the inferior colliculus, the medial geniculate nucleus and the auditory cortex. The results show that the lateral lemniscus projects to the central nucleus of the inferior colliculus which in turn projects principally to the ventral division of the medial geniculate nucleus but to a lesser extent to the magnocellular division of the medial geniculate nucleus. The final step in the pathway to the cortex is achieved by a projection from the ventral division to the fourth layer of auditory koniocortex. There appear to be several auditory pathways parallel to this primary path. The lateral lemniscus projects to the dorsal division of the medial geniculate nucleus; the deeper layers of the superior colliculus project to the posterior nucleus; and both the dorsal division and the posterior nucleus project to the belt caudal to auditory koniocortex. The caudal division of the medial geniculate nucleus may constitute a relay in still another path from the pericentral division of the inferior colliculus. Finally, the magnocellular division also appears to be distinct insofar as its cortical projections are confined chiefly to the deeper layers. A comparison between the tree shrew and the cat reveals a similar organization in the two species. In the cat the starting point for understanding the organization of the several auditory pathways is the distinction between a core cortical zone which corresponds to koniocortex and to AI and a peripheral belt. The core receives essential projections from the ventral division; the belt receives sustaining projections from the cell groups which surround the ventral division. It is reasonable to hypothesize that this difference between the core and the belt is characteristic of all mammals.     

The serotoninergic fibers in the dorsal lateral geniculate nucleus of the cat: distribution and synaptic connections demonstrated with immunocytochemistry

The distribution, morphology, and synaptic contacts of serotoninergic fibers were studied with immunocytochemical methods in the lateral geniculate complex of the cat. The serotonin-immunoreactive fibers are diffusely distributed throughout the main laminae of the dorsal lateral geniculate nucleus (dLGN) and the perigeniculate nucleus (PGN) and reach a particular density in the ventral lateral geniculate nucleus (vLGN). The labeled fibers are in most cases very thin and sometimes varicose. There is no obvious order in their distribution pattern except that they sometimes partially encircle the unlabeled cell bodies of the dLGN. The synaptic connections of the serotoninergic fibers were investigated mainly in the A laminae of the dLGN. Few synaptic complexes were found, most of them with asymmetric morphology. The postsynaptic elements were small dendritic profiles. Perisomatic serotoninergic fibers were seen, but no convincing synaptic contacts were found between labeled fibers and cell somata. In the dLGN, serotoninergic profiles were almost exclusively confined to the extraglomerular neuropile. In the PGN serotoninergic fibers also contacted dendritic profiles and formed asymmetrical synapses, but as in the geniculate, synaptic specializations were very rare.     

Genesis of neurons in the dorsal lateral geniculate nucleus of the cat

Neurons that will ultimately form the dorsal and ventral lateral geniculate nuclei, the medial interlaminar nucleus, the perigeniculate nucleus, and the nucleus reticularis of the cat undergo their final cell division beginning on, or slightly before, embryonic day 22 (E22) and ending on, or before, E32. Early in this period, neurogenesis proceeds for all of these geniculate nuclei, whereas only in the dorsal lateral geniculate nucleus does cell birth continue until E32. Distinct spatiotemporal gradients of cell birth are not obvious within any of the individual geniculate nuclei. For the dorsal lateral geniculate nucleus in particular, and for the other geniculate nuclei in general, neurons born early in this period exhibit a full range of adult soma sizes, including large and small neurons. Neurons born late in this period exhibit only small adult somas. The location and size of a neuron within the dorsal lateral geniculate nucleus provide clues to that cell's functional properties. On the basis of presently available information regarding the relationship between structure and function of neurons in the cat's dorsal lateral geniculate nucleus, the findings described here suggest that all functional classes of neurons in the dorsal lateral geniculate nucleus are born at the same time throughout most of this period.     

GABAergic circuitry in the dorsal division of the cat medial geniculate nucleus

Gamma-aminobutyric acid (GABA) is the main inhibitory neurotransmitter of the thalamus. We used postembedding immunocytochemistry to examine the synaptic organization of GABA-positive profiles in the dorsal superficial subdivision of the cat medial geniculate nucleus (MGN). Three groups of GABA-positive profiles participate in synapses: axon terminals, dendrites, and presynaptic dendrites. The presynaptic GABA-positive terminals target mainly GABA-negative dendrites. The GABA-positive postsynaptic profiles receive input primarily from GABA-negative axons. The results indicate that the synaptic organization of GABA-positive profiles in the dorsal superficial subdivision of the MGN nucleus is very similar to that in other thalamic nuclei.     

Qualitative and quantitative analyses of the patterns of retinal input to neurons in the dorsal lateral geniculate nucleus of the cat

In the visual system of the cat the projection from the retina to the lateral geniculate nucleus has been studied extensively. However, the patterns of synaptic contacts made by individual axons onto individual cells have not been described. In this study these patterns have been examined for class 1 cells (Guillery: J. Comp. Neurol. 128:21, '66). Retinogeniculate axons and lateral geniculate neurons are labeled with horseradish peroxidase (HRP) via injections into the optic tracts and optic radiations, respectively. Sections are then processed for combined light and electron microscopic analysis. They are examined with the light microscope to identify labeled lateral geniculate neurons that appear to be contacted by labeled retinal axons. These cells and axons are then analyzed by a computerized microscope system, and sites of apparent synaptic contact are recorded. This light microscopic analysis indicates that individual class 1 cells are contacted by many retinogeniculate axons (> 10) and that each of these axons contacts many lateral geniculate neurons (> 20). Some axons make numerous contacts that are concentrated onto a few dendrites, while others make only a few contacts, which are spread over several dendrites. In all cases, the majority of contacts are on the dendritic shafts of relatively thick secondary and tertiary dendrites. Electron microscopic analysis confirms that most of the contacts identified with the light mciroscope are synaptic. It also reveals that labeled and unlabeled retinal axons can innervate the same dendritic segment. Finally, one cell was studied that had its soma and most of its dendrites in lamina A1 but some of its dendrites extended into lamina A. This cell received input from retinal axons in both layers, thus suggesting that it may have been binocularly excitable. © 1993 Wiley-Liss, Inc.     

Mode of termination of afferents from the thalamic reticular nucleus in the dorsal lateral geniculate nucleus of the rat

The terminals of axons projecting to the dorsal lateral geniculate nucleus from the thalamic reticular nucleus were identified by electron microscopy 8–24 h after placing small lesions in the ipsilateral reticular nucleus. The terminals contained flattened synaptic vesicles and made Gray type II axo-dendritic synaptic contacts with geniculate neurons. Their identification as F-axons accords well with physiological evidence for a powerful monosynaptic inhibitory input to geniculocortical projection cells from reticular nucleus neurons.     

Development of the dorsal lateral geniculate nucleus in normal and visually deprived cats

Although much is known about the cell size changes that take place in the cat dorsal lateral geniculate nucleus as a result of visual deprivation, very little is known about the time course of either of these changes or the changes that occur during normal development. In addition, all previous studies of lateral geniculate nucleus cell size have been confined to the dorsal laminae A and A1 since the more ventral “C” laminae are impossible to identify in normal Nissl stained material. However, it is possible to extend the cell measurement data to laminae C, C1, and C2 by using autoradiographic techniques. Cross-sectional area measurements of dorsal lateral geniculate nucleus cells were made in 47 normally reared kittens and 45 monocularly deprived kittens. All of the normal kittens and 39 of the 45 deprived kittens were studied during the first 70 days of postnatal life. Six deprived cats used to study the deprivatin induced changes in cell size in the “C” laminae were allowed to survive for longer periods. In normal kittens, lateral geniculate nucleus cells grow rapidly during the first four weeks of life. Cells in the deprived layers also grow rapidly during this time, however, at the end of the first month their growth stops and a slow shrinkage takes place over the next several weeks. In the ‘C’ laminae of deprived cats significant changes in cell size are confined to layer C. Although many of the deprived cats show greater deprivation induced changes in cell size in the binocular segment of the lateral geniculate nucleus than in the monocular segment, other cats show approximately equal changes in cell size in the two regions. In addition, some cats exhibit little, if any, deprivation induced change in lamina A cell size but do show quite severe cell shrinkage in lamina A1.     

Genesis of morphologically identified neurons in the dorsal lateral geniculate nucleus of the cat

Neurons in the dorsal lateral geniculate nucleus of the cat can be grouped into five morphological classes based on a variety of structural characteristics. These same structural characteristics can serve as morphological signatures for the three physiological classes (X, Y, W) of neurons found in this nucleus. The purpose of this study was to determine if a relationship exists between the birthdate of neurons within the dorsal lateral geniculate nucleus and the adult morphology of those neurons. Seven cats, each of which had received a single injection of 3H-thymidine, were studied. A total of 2,138 Golgi-impregnated neurons were identified in the dorsal lateral geniculate nuclei of these seven cats; 1,517 of these neurons were successfully resectioned and recovered, of which 385 (25%) were found to contain the 3H label. Neurons from each of the five morphological classes were labeled in each of the six animals that received a 3H-thymidine injection between embryonic day 24 (E24) and E28. Class 3 and class 5 neurons were labeled in a cat injected with 3H-thymidine on E30. These findings demonstrate that the development of the morphological class of a neuron in the dorsal lateral geniculate nucleus is independent of the time of its final cell division. Further, given the relationship that exists in the cat's dorsal lateral geniculate nucleus between neuronal structure and function, the present findings suggest that the different physiological classes of cells found in this nucleus undergo their final cell divisions throughout most of the period of neurogenesis except that the functional role of neurons born late in this period may be more restricted.     

The fine structure of the perigeniculate nucleus in the cat

The fine structure of the cat's perigeniculate nucleus has been analyzed and compared to that of dorsal thalamic relay nuclei. Golgi preparations and electron micrographs of perigeniculate cells commonly show somatic spines. The most common presynaptic elements for these spines and for the adjacent perikaryal surfaces are relatively large axon terminals containing round synaptic vesicles and making multiple asymmetric contacts. These "RLD" terminals (so termed for their round vesicles, large average size of the terminals, and dark mitochondria) are also presynaptic to dendritic spines and shafts of proximal and secondary dendrites. Comparisons with adjacent parts of the dorsal lateral geniculate nucleus show that these RLD terminals are cytologically distinct from retinogeniculate terminals and that small numbers of RLD terminals also occur in the geniculate A laminae. Three other major classes of perigeniculate synaptic terminals, resemble major classes of terminals in the dorsal lateral geniculate nucleus. These include two types of terminal with flat or ovoid synaptic vesicles and dark mitochondria, "FD1" and "FD2" terminals, and a class of small terminal with densely clustered round vesicles and dark mitochondria, "RSD" terminals. RSD terminals, which resemble corticogeniculate axon terminals, represent the only class of perigeniculate terminal that does not contact perikarya. FD2 terminals resemble lateral geniculate presynaptic dendrites and participate in serial and triadic synaptic contacts, being both pre- and postsynaptic; however, in contrast to the arrangement characteristic of thalamic relay nuclei, these contacts do not occur within synaptic glomeruli. A fifth major class of perigeniculate presynaptic terminal has large flat or polymorphic synaptic vesicles and pale mitochondria. These "FP" terminals are seen infrequently in the lateral geniculate A laminae. Similarities between perigeniculate and lateral geniculate fine structure may relate in part to common sources of afferent input to the two nuclei.     

Quantitative comparison of retinal synapses in the dorsal and ventral (parvicellular) C laminae of the cat dorsal lateral geniculate nucleus

Three physiological classes of retinal ganglion cell project to the cat dorsal lateral geniculate nucleus (DLGN). The dorsal laminae A, A1, and magnocellular C receive X and Y retinal input, whereas the ventral parvicellular laminae C1 and C2 receive predominantly W input. We have compared quantitatively the retinal synaptic terminals of the dorsal and ventral laminae to determine whether there are morphological differences in the terminals that correspond to their different response properties. Anterogradely labeled retinal synaptic terminals in all laminae contained pale mitochondria and large, round synaptic vesicles. However, retinal terminals with pale mitochondria varied in size and synaptic organization in different laminae. The terminals in the A laminae were, on average, quite large and made numerous contacts with conventional dendritic profiles and with profiles that themselves contained synaptic vesicles (F2 profiles). The terminals in lamina C that contained pale mitochondria had a smaller overall mean area. Terminals with pale mitochondria in C1 and C2 were almost all small and synapsed with F2 profiles less frequently than did terminals in the A laminae or in lamina C. These results provide quantitative evidence that visual areas receiving W-type retinal input contain smaller retinal terminals and have a different synaptic organization from that of laminae receiving X and Y input.     

Synaptic organization of inhibitory pathways to principal cells in the lateral geniculate nucleus of the cat

The synaptic organization of inhibitory systems in the lateral geniculate nucleus of the cat was studied with intracellular recording techniques. Principal cells of both X- and Y-type received disynaptic inhibition of the feed-forward type from retinal ganglion cells, in most cases from both eyes. All cells also received inhibition of the recurrent type via a subcortical pathway, as postulated in many earlier studies.     

Ultrastructure of cholinergic synaptic terminals in the thalamic anteroventral,ventroposterior, and dorsal lateral geniculate nuclei of the rat

The principal relay nuclei of the thalamus receive their cholinergic innervation from two midbrain cholinergic groups: the pedunculopontine tegmental nucleus and the laterodorsal tegmental nucleus. The different thalamic nuclei exhibit populations of cholinergic axons which vary in density and morphology when examined at the light microscopic level. However, the ultrastructure of the cholinergic terminals in different thalamic nuclei has not been described. This study was undertaken to confirm that synaptic contacts are formed by cholinergic axons in several principal thalamic relay nuclei, to describe their ultrastructural morphology, and to identify the types of postsynaptic elements contacted by cholinergic synaptic terminals. The thalamic nuclei examined in this study are the dorsal lateral geniculate nucleus, ventroposteromedial nucleus, ventroposterolateral nucleus, and anteroventral nucleus. Our results confirm that cholinergic axons form synaptic terminals in these thalamic nuclei. Cholinergic synaptic terminals contact structures outside the characteristic synaptic glomeruli, are never postsynaptic, and have morphologies and postsynaptic targets which differ among the thalamic nuclei. In the ventroposterior nuclei, cholinergic terminals form asymmetric synaptic contacts onto larger dendrites in the extraglomerular neuropil. In the anteroventral nucleus, cholinergic terminals form both symmetric and asymmetric synaptic contacts onto dendrites and somata. Cholinergic terminals in the anteroventral nucleus are larger than those in other nuclei. In the dorsal lateral geniculate nucleus, cholinergic terminals contact both somata and dendrites in the extraglomerular neuropil, but the synaptic contacts in this nucleus are symmetric in morphology.     

Ventral lateral geniculate nucleus projects to the dorsal lateral geniculate nucleus in the cat

The lamina C3 of the dorsal lateral geniculate nucleus of the cat does not receive retinal projections but instead receives visual information from the small subpopulation of W-type ganglion cells via the upper substratum of the stratum griseum superficiale of the superior colliculus. We herein report a projection from the lateral division of the ventral lateral geniculate nucleus into the lamina C3 of the dorsal lateral geniculate nucleus. As the lateral division receives projections from the contralateral retina and the ipsilateral upper stratum griseum superficiale of the superior colliculus, we suggest that these regions make up a small cell type W-cell neuronal network that provides visual information to layer I of the striate cortex via the lamina C3.