A multimodal pathway including the basal ganglia in the feline brain.

The purpose of this paper is to give an overview of our present knowledge about the feline tecto-thalamo-basal ganglia cortical sensory pathway. We reviewed morphological and electrophysiological studies of the cortical areas, located in ventral bank of the anterior ectosylvian sulcus as well as the region of the insular cortex, the suprageniculate nucleus of the thalamus, caudate nucleus, and the substantia nigra. Microelectrode studies revealed common receptive field properties in all these structures. The receptive fields were extremely large and multisensory, with pronounced sensitivity to motion of visual stimuli. They often demonstrated directional and velocity selectivity. Preference for small visual stimuli was also a frequent finding. However, orientation sensitivity was absent. It became obvious that the structures of the investigated sensory loop exhibit a unique kind of information processing, not found anywhere else in the feline visual system.

Neuroanatomical study on the tecto-suprageniculate-dorsal auditory cortex pathway in the rat.

Previous anatomical and physiological studies suggest that the superior colliculus sends integrated sensory information to the multimodal cortical areas via the thalamic suprageniculate nucleus (SG). However, the detailed distribution of rat tecto-SG axon terminals and SG neurons projecting to the multimodal cortex, as well as synaptic connections between these tectal axons and SG neurons, remains unclear. In this study, the organization of the tecto-thalamo-cortical pathway was investigated via combined injections of anterograde and retrograde tracers followed by light and electron microscopic observations. Injections of a retrograde tracer, cholera toxin B subunit (CTB), into the temporal cortex, area 2, dorsal part (Te2D), and injections of an anterograde tracer, biotinylated dextran amine (BDA), into the deep layers of the superior colliculus produced the following results: (1) Retrogradely CTB-labeled neurons were found throughout SG, predominantly in its rostral part. CTB-labeled neurons were also found in other cortical areas such as the visual cortex, the auditory cortex, the parietal association cortex, and the perirhinal cortex. (2) Anterogradely BDA-labeled axons and their terminals were also observed throughout SG. Dual visualization of BDA and CTB showed that retrogradely labeled SG neurons and anterogradely labeled tectal axon terminal boutons overlapped considerably in the rostral part of SG, and their direct synaptic contacts were also confirmed via electron microscopy. These findings suggest that multimodal information from the superior colliculus can be processed directly in SG neurons projecting to Te2D.

Overlap of nigrothalamic terminals and thalamostriatal neurons in the feline lateralis medialis-suprageniculate nucleus.

The lateralis medialis-suprageniculate nucleus (LM-Sg) of the feline posterior thalamus is a relay nucleus with a clear visuomotor function. In this study, we examined the distribution of axon terminals of the nigral afferent to the LM-Sg following injection of an anterograde tracer, biocytin, into the substantia nigra pars reticulata, and the distribution of the thalamostriatal projection neurons in the LM-Sg following the injection of wheat germ agglutinin conjugated with horseradish peroxidase (WGA-HRP) as a retrograde tracer into the caudate nucleus. The biocytin-labeled terminal-like puncta were located in the ventromedial portion of this nucleus in such a way that most of the labeled elements took the form of swellings having boutons in places, while a minority appeared in clusters of 3-5 large terminal-like puncta. The retrograde WGA-HRP-labeled neurons were also found in the ventromedial part of the LM-Sg, and the distributions of labeled nigrothalamic axon terminals and labeled thalamostriatal projection neurons therefore overlapped in this region. The present results indicate that the nigral afferent may make synaptic contacts directly with the thalamostriatal projection neurons within the LM-Sg.

Postnatal development of the corticothalamic projections from the visual association cortex to the lateralis posterior complex in the cat.

The postnatal development of the corticothalamic projection from the lateral suprasylvian cortex (LS) to the lateral medialis-suprageniculate nucleus (LM-Sg) of the cat thalamus was assessed by means of the anterograde tracer biocytin. In the adult, two types of corticothalamic fibers were found: type I established a network of fine fibers present throughout the LM-Sg, it was characterized by a linear sequence of small (less than 0.5 microm in diameter), single terminal boutons making contact mainly with thin dendrites and/or dendritic spines. Type II, found less frequently, gave off short, side branches near axon terminals and formed clusters of 5-10 large terminal boutons (0.5-1.5 microm in diameter), making contact predominately with medium-sized dendrites and/or vesicle-containing profiles, forming a synaptic glomerulus. At birth (P0), anterogradely-labeled fibers were found in the LM-Sg as in adults. In the early postnatal period (until P6) as well as around the time of eye-opening (P7-P10) to P21, neonatal fibers were largely unbranched many of them having axons tipped with growth cones. Axon terminals containing synaptic vesicles were rarely observed but when present these exhibited considerable variation in their morphological appearance of synapses. Thus, it was not possible to categorize them into the two types of axons which characterize the adult. After P25, terminal swellings bearing a close resemblance to those of type II fibers begin to appear. In this way, the main two corticothalamic fiber types could be identified. These findings demonstrate that significant postnatal changes occur in the synaptology of corticothalamic fibers in the LM-Sg, particularly with the maturation of type II fibers.

Visual receptive field properties of excitatory neurons in the substantia nigra.

The substantia nigra has been widely regarded as a structure involved in visuomotor co-ordination, but little is known about the sensory background of its function. Here we give a detailed description of the visual receptive field properties of excitatory substantia nigra neurons. The visual responses of 59 excitatory neurons were recorded in both the substantia nigra pars reticularis and the pars compacta of halothane-anesthetized, immobilized, artificially respirated cats. The substantia nigra neurons were not responsive or exhibited very low sensitivity to stationary visual stimulation. The units responded optimally to small stimuli moving at intermediate or high velocities in their extremely large receptive field. We observed no signs of retinotopical organization within the substantia nigra. A majority of the units exhibited narrow direction tuning and high direction selectivity, while a smaller proportion of them were broadly tuned and not direction-sensitive. Our results suggest that the visual properties of the excitatory substantia nigra units are quite similar to those of the superior colliculus and other extrastriatal structures that receive tectal afferents. This supports the notion that the substantia nigra processes dynamic visual information and that its excitatory visual neurons are modulated by the extrageniculate tectal visual system of the mammalian brain.

Postnatal development of the retinal projection to the nucleus of the optic tract and accessory optic nuclei in the hooded rat.

Retinal projections to the nucleus of the optic tract (NOT) and accessory optic nuclei (AON) were studied in the postnatal hooded rat after monocular injection of cholera toxin B subunit (CTB) into the vitreous chamber of the eye. At all postnatal ages, retinal axons were labeled sensitively; they revealed dense projections to the contralateral, and sparse but distinct projections to the ipsilateral, NOT and AON. The CTB labeling enabled the first delineation of the complete morphology of developing retinal axons in the ipsilateral NOT and AON. From postnatal day (P) 1 to P3, axons with complex growth cones were seen, and unbranched collaterals with simple growth cones increased and extended gradually. At P6, complex growth cones disappeared while branched collaterals with simple growth cones as well as small-sized varicosities increased. By P12 (two days before eye-opening) the adult-like pattern of terminal arbors appeared. The branched collaterals with tiny, small-sized varicosities present probably represented developing synaptic boutons. At P16 (after eye opening), the pattern of terminal arbors was well developed, almost to the same extent as in the adult. By contrast, a broadly distributed, transient retinal projection around NOT and AON was gradually eliminated; it started to disappear during the first few postnatal days, and was fully retracted by the time of eye-opening time to a pattern normal for the adult.

Immunocytochemical study of G(i)2alpha and G(o)alpha on the epithelium surface of the rat vomeronasal organ.

To investigate in detail the distribution of G protein subtypes G(i)2alpha and G(o)alpha along the surface of the vomeronasal epithelium, we used double labeling immunocytochemical methods and electron microscopy. We examined the immunoreactivity of these surface structures with antibodies against G(i)2alpha and G(o)alpha. G(i)2alpha- and G(o)alpha-positive cells were observed at the epithelial surface and were evenly distributed. Electron microscopy revealed that strong immunoreactivities to both antibodies were observed on the microvilli and knob-like surface structures of receptor cells. No immunoreactivity was found on the microvilli or surface membranes of supporting cells. This expression pattern is similar to that reported for putative pheromone receptors. These data confirm that there are two distinct classes of vomeronasal receptor cells expressed at the surface of the epithelium. These two classes of receptors correspond to the same G(i)2alpha- and G(o)alpha-positive cells distributed in cell body layers of the epithelium and in the axon terminals in the accessory olfactory bulb.

Ultrastructural organization of transmitters in the cat lateralis medialis-suprageniculate nucleus of the thalamus: an immunohistochemical study.

The lateralis medialis-suprageniculate nuclear (LM-Sg) complex of the cat's posterior thalamus receives a rather wide variety of inputs from diverse cortical and subcortical areas. Previous ultrastructural studies of this nucleus demonstrated the presence of four types of vesicle-containing profiles and characterized some of these as gamma-aminobutyric acid (GABA)-containing terminals (Norita and Katoh [1987] J. Comp. Neurol. 263:54-67; Norita and Katoh [1988] Prog. Brain Res. 75:109-118). The present study has extended these observations by examining the immunoreactivity (ir) of LM-Sg, with antibodies raised against aspartate (Asp), glutamate (Glu), GABA, the acetylcholine (ACh) marker, choline acetyltransferase (ChAT), and substance P (SP), by using light and electron microscopy. Neuronal somata immunopositive for the excitatory amino acids (EAAs) Asp and Glu, were of medium size. EAA-ir terminals also were of medium size and contained round synaptic vesicles; they made asymmetrical synaptic contacts with dendritic profiles. Neuronal somata immunopositive for GABA were small. GABA-positive terminals also were small and contained pleomorphic synaptic vesicles; they formed symmetrical synaptic contacts with dendritic profiles. No neurons immunolabeled for ChAT were found. Terminals immunopositive for ChAT were small and contained round synaptic vesicles; these made symmetrical synaptic contacts, asymmetrical synaptic contacts, or both, of the en passant type with dendritic profiles. SP-immunolabeled neuronal somata were not found. Immunolabeled terminals were small, contained round synaptic vesicles, and made asymmetrical synaptic contacts with dendritic profiles. ChAT-ir and SP-ir axon terminals were not expressed evenly within LM-Sg. This difference in distribution suggests that within the LM-Sg, there may be a difference in specific sensory processing functions which correlate with transmitter type.

Distribution of terminals from pedunculopontine tegmental nucleus and synaptic organization in lateralis medialis-suprageniculate nucleus of cat's thalamus: anterograde tracing, immunohistochemical studies, and quantitative analysis.

The cat's lateralis medialis-suprageniculate nuclear complex (LM-Sg) in the thalamus receives input from various brain regions such as the superior colliculus, brain stem, and spinal cord, as well as from visual association cortex. In a previous study, we demonstrated that LM-Sg receives cholinergic fibers from the pedunculopontine tegmental nucleus (PPT) and that cholinergic terminals make synaptic contacts with the dendrites of glutamatergic projection neurons and of GABAergic interneurons (Hoshino et al., 1997). In this study, we investigate the distribution and the organization of PPT terminals by means of a combined anterograde tracer (biotinylated dextran amine, BDA) and immunohistochemical methods. When stained by acetylcholinesterase (AChE), the LM-Sg is not uniformly immunoreactive, but rather is patchily labeled and shows a streaming type of reactivity. The tissue content appears high in enzyme activity in AChE-positive zones and is much lighter in activity in AChE-negative zones. We compared the synaptic organization between AChE-positive and AChE-negative portions of the LM-Sg in separate groups of electron-microscopic material: four types of vesicle containing profiles (RS, RL, F1, and PSD) as well as synaptic glomeruli were observed in this nucleus. Among these, the PSD profiles were observed more frequently in AChE-positive portions than in AChE-negative zones. Furthermore, the number of glomeruli was significantly higher in AChE-positive than in AChE-negative zones. Following the injection of BDA into PPT, labeled terminals within LM-Sg were rather more concentrated in the AChE-positive portion. Although the majority of PPT terminals made synaptic contacts with dendrites in the neuropil, a few terminals were involved in the synaptic glomeruli. The present results show that the synaptic organization is distinctly different between the AChE-positive and AChE-negative portions of LM-Sg. These results suggest that the AChE-positive portions of LM-Sg are relatively more involved in integrating information arising from a diverse set of inputs and processing that information within glomeruli in a complex manner than occurs in the AChE-negative portion of LM-Sg.

Metabotropic glutamate receptor 2/3 immunoreactivity in the developing rat cerebellar cortex.

In adult rat cerebellar cortex, the metabotropic glutamate receptors (mGluRs) 2 and 3 (mGluR2/3) are present in somata, dendrites, and terminals of Golgi cells as well as in presumed glial processes (Ohishi et al. [1994], Neuron 13:55-66). In the present study, spatiotemporal changes in immunostaining for mGluR2/3 were examined in postnatal rat cerebellar cortex. mGluR2/3-immunoreactive Golgi cell somata appeared first in the internal granular layer at postnatal day 3 (P3) and were restricted to lobules IX and X; however, by P5, they were present in all lobules. Immunoreactive Golgi cell axons were adult-like, appearing as tortuous fibers with clusters of varicosities. They were observed first in the internal granular layer at P7 and increased in number and complexity with time. It was confirmed that mGluR2/3-immunoreactive Golgi cell axon terminals belong to the synaptic glomerulus by P10. Immunoreactive Golgi cell dendrites extending into the molecular layer became prominent after P15. By that time, the immunostaining pattern was characteristic of Golgi cells, as seen typically in adults. Many intensely immunoreactive radial processes existed at birth (P0). These traversed the molecular and external granular layers, reaching the pial surface in every cerebellar lobule. Because they showed coimmunoreactivity for glial fibrillary acidic protein, they were confirmed to be Bergmann glial fibers. After P9, they began to lose immunoreactivity at the portion corresponding to the molecular layer, while an immunostained granular pattern appeared in that layer. Immunoreactive radial processes, however, remained in the external granular layer, and finally, at P21, they disappeared together along with the external granular layer. Granular staining in the molecular layer reached background levels at this time. These spatiotemporal changes in mGluR2/3 distribution suggested that there may be distinct roles for mGluR2/3 in Golgi cells and Bergmann glial cells during the early postnatal period. mGluR2/3 in Golgi cells might be associated closely with systemic maturation, whereas mGluR2/3 in Bergmann glia might be needed for neuron-glia interactions related to granule cell development.

A light and electron microscopic analysis of the convergent retinal and visual cortical projections to the nucleus of the optic tract (NOT) in the pigmented rat.

It is generally known that the nucleus of the optic tract (NOT) subserves visuomotor relations between the retina and preoculomotor structures as the only subcortical pathway mediating optokinetic responses (OKR) in mammals. We have examined the projections from the retina and visual cortical areas (areas 17, 18a and 18b) to NOT using tracers (wheat germ agglutinin-conjugated horseradish peroxidase, WGA-HRP and cholera toxin B subunit, CTB) in order to clarify how these two different functional inputs to OKR are organized. CTB injection into the vitreous body resulted in anterograde label almost exclusively in the contralateral NOT. Ultrastructually, the size of the retinal axon terminals was small (up to 0.7 micron in diameter), contained round synaptic vesicles and pale mitochondria, and made asymmetrical synaptic contacts with both GABA-positive and GABA-negative NOT neurons. Visual cortical area 17 and the transitional area between area 17 and 18a, or between area 17 and 18b projected their axons to the ipsilateral NOT. Ultrastructually, the size of the cortical axon terminals was small (up to 0.5 micron in diameter), contained round synaptic vesicles, and made asymmetrical synaptic contacts only with GABAnegative NOT neurons. With light and electron microscopical observation, there was a considerable overlap in the cortico-NOT and retino-NOT projection pattern: GABA-negative (presumably NOT projection) neurons simultaneously receive input from both cortical and retinal terminals. From these results, it seems reasonable to postulate that inputs from visual cortical areas in the pigmented rat cooperate with those from the retina in controlling OKR.

The organization of the thalamostriatal projection from the lateral posterior thalamic nuclear complex (LP) in the pigmented rat.

We investigated the thalamostriatal projection of the rat using biotinylated dextran amine (BDA) and wheat-germ agglutinin horseradish peroxidase (WGA-HRP). To obtain the patch/matrix compartments of the striatum (ST), we used mu-opioid receptor (MOR) immunoreaction labeling. Thus, an MOR-positive 'patch' was indicated by a darkly stained spot, while the MOR-negative 'matrix' was displayed as a non-immunoreactive region. A small injection of BDA was made in a subregion of the lateral posterior thalamic nucleus (LP). The LP-ST fibers originated in all subregions of LP and terminated in the dorsocaudal portion of ST, where the corticostriatal fibers from the visual cortex terminate (Serizawa et al. 1994). These LP-ST fibers and terminals were concentrated in the MOR-negative matrix compartment. Electron microscopic observations showed that the LP-ST terminals made asymmetrical synaptic contacts mainly (70%, n = 30) with the dendritic spines of the presumptive ST-output neurons, and fewer (30%) contacted dendritic shafts. The present results provide anatomical support for the contention that ST-output spiny neurons of the matrix that project to the pars reticulata of the substantia nigra or globus pallidus, may be influenced directly by the LP-ST projection.

Laminar organization of the intrinsic connection neurons and their axon terminals in the lateral suprasylvian cortex of the cat.

The lateral suprasylvian cortex (LS) of the cat has numerous interconnections with other visual cortical areas, as well as with subcortical structures implicated in visually-guided behavior. However, little data are available regarding connections within the LS itself. In order to investigate the intrinsic connections within LS, we have examined the laminar patterns of terminal and cellular labeling following injections of either the anterograde tracer biocytin, or the retrograde tracer, WGA-HRP, into the subregions of the LS (AMLS, PMLS, ALLS, and PLLS). Tracer deposits spanning all cortical layers in a LS subregion basically resulted in labeling of axons and terminals (biocytin cases) or cells of origin (WGA-HRP cases) in all layers, although intensity of the labeling seemed to be different among subregions. The present study demonstrated that the interconnections among LS subregions provide no clear evidence of simple hierarchial relationships between regions.

Normal ontogenic observations on the expression of Eph receptor tyrosine kinase, Cek8, in chick embryos.

The spatio-temporal pattern of expression for the Eph receptor tyrosine kinase, Cek8, was observed in normal chick embryos from H-H stage 6 to 23 by immunohistochemical techniques. Expression of Cek8 was already present in embryos at H-H stage 6, where it was located in the neural plate of the brain region, paraxial mesoderm, and the primitive streak. Regions expressing Cek8 subsequently increased during development to include the neural folds of the brain, rhombomeres 3 and 5, the caudal part of the neural plate, neural crest cells related to the formation of glossopharyngeal nerve ganglia, invaginated cells throughout the primitive groove and the epithelium of the rudiment of the gall bladder. Cek8 was also expressed in the mesenchymal cells of the pharyngeal arches, allantoic stalk and limb buds as well as in the areas surrounding the eye vesicles and nasal pits. Furthermore, cells in the tail bud progressing to the secondary neurulation expressed Cek8. Thus, the spatiotemporal patterns of Cek8 expression appears to have intimate relationships with tissue rebuilding, the maturation of differentiated cells, and the spatial organization of tissues. Consequently, it appears that Cek8 plays an integral role in the developmental events leading to the formation of a wide--though not inclusive--variety of tissues and organ systems.

Early development of the oligodendrocyte in the embryonic chick metencephalon.

It has been demonstrated that the spinal cord oligodendrocytes in the vertebrates arise in the ventral ventricular zone adjacent to the floor plate in their early development. Because of the similarities of basic structures in the spinal cord and metencephalon, it is probable that the mode of early oligodendrocyte development in the metencephalon is the same as that in the spinal cord. We examined this possibility in chick embryos, using monoclonal antibodies O1 and O4, markers for oligodendrocyte lineage. An O4-positive (O4+) cell focus was observed in the medial ventricular zone of E5 chick ventral metencephalon (the earliest stage examined), adjacent to the floor plate. At E6, O4+ cells were dispersed from the medial to the lateral pons and, at E7, to the cerebellar anlagen. O4+ cells in the E6 brainstem and in the E7 cerebellum were unipolar in shape, whereas one day later, some of the labeled cells were multipolar with a few thin processes. O1+ oligodendrocytes first appeared at E8 in the ventromedial part of the pons and were distributed throughout the pons at E10 and in the cerebellum at E12. Explants from three subdivisions of the metencephalon (medial and lateral pons, and cerebellum) from E5 to E8 chick embryos were separately cultured to confirm the potential for generation of oligodendrocyte lineage. O4+ cells appeared in the culture of the E5 medial pons (the earliest stage examined), in the E6 lateral pons, and in the E7 cerebellum. In addition, E7 was the youngest stage from which cerebellar explants were able to generate O1+ oligodendrocytes. Our results clearly demonstrated the in vivo morphology of oligodendrocyte precursors in the metencephalon and their developmental appearance in a ventral-to-dorsal manner. From the bipolar morphology of O4+ cells and the spacio-temporal continuity of the dispersion, it is inferred that the initial dispersion of O4+ cells may involve oligodendrocyte migration from the focus of the medial pons to the lateral and dorsal parts of the metencephalon.

Cholinergic innervation of the lateralis medialis-suprageniculate nuclear complex (LM-Sg) of the cat's thalamus: a double labeling immunohistochemical study.

The purpose of this study was to investigate morphological characteristics of the synaptic relations of choline acetyltransferase (ChAT)-positive terminals that are made with a variety of post-synaptic profiles in the lateralis medialis-suprageniculate nuclear complex (LM-Sg) using ChAT, gamma-aminobutyric acid (GABA) and glutamate immunohistochemistry in combination with electron microscopical observations. The ChAT immunopositive profiles make asymmetrical synaptic contacts with glutamate immunopositive dendrites that are presumably derived from projection neurons, and/or GABA immunopositive interneurons. The present results indicate that ascending cholinergic mechanisms may be important for modifying information in both the extrinsic and intrinsic circuitries of LM-Sg.

Morphological analysis of the early development of the chick neural tube separated from the floor plate and notochord.

When the neural tube of avian embryos is separated from the notochord and floor plate, motoneurons in the spinal cord fail to develop. In order to investigate the factors involved in this phenomenon, cell proliferation activity and cell death were observed following paramedian incision of the neural tube at the level of the segmental plate using colchicine, BrdU, and TUNEL methods. If the notochord and/or floor plate produces a substance(s) that promotes cell division in the basal plate neuroepithelium or that supports the survival of the motoneuron's neuroblasts, mitotic figures should not be present in the neuroepithelium nor should substantial cell death be observed in the ventral aspect of the notochord- and floor plate-deprived neural tube. Surprisingly, however, neither result was observed in the present experiments, with the exception of a considerable amount of homogeneously distributed cell death. Neuroepithelial cells continued to proliferate and gave rise to neuroblasts. Nevertheless, motoneurons failed to develop, and the neural tube was enveloped by only the basement membrane of the alar plate (S. Hirano and H. Tanaka, 1994, Dev. Growth Differ: 36, 481-488). These morphological results revealed that the cause of the development of the anterior horn lacking a neural tube in the notochord- and neural tube-eliminated embryos is not the elimination of the source of the surviving factor(s) of the motoneuron's neuroblasts, but rather the elimination of the signals to induce the motoneurons, derived from the notochord and/or floor plate. The larger amount of cell death in the neural tube on the experimental side suggests that a nonspecific survival factor(s), necessary for the survival of a variety of types of neuroblasts, is also produced by the notochord and/or floor plate.

Extrinsic and intrinsic connections of the cat's lateral suprasylvian visual area.

The lateral suprasylvian visual area (LS) is known to have numerous interconnections with visual cortical areas as well as with subcortical structures implicated in visually-guided behaviors. In contrast, little data is available regarding connections within the LS itself. In order to obtain information about intra-areal connections and to re-investigate LS connectivity with various cortical and subcortical areas, the traces (biocytin or WGA-HRP) was injected into various loci along the medial and lateral banks of the LS. The anterograde tracer, biocytin injections into both medial and lateral bank produced label contained within the respective bank that extended rostrally and caudally from the infection site. In addition, following medical bank injections, considerable label was distributed throughout the fundus and, to a lesser extent, in the lateral bank. In contrast, no label could be detected in the medial bank after lateral bank injections, and, although label was observed in the fundus, it was restricted to the most lateral aspects. Moderate labeling could be observed in the medial bank following the tracer injection into the most rostral aspect of the lateral bank. It is likely that input derived from various visual cortical areas which project to the medial bank of the LS has access to this intra-areal circuitry. This may provide a route by which visual cortical information can be relayed to other cortical and subcortical structures involved in visually-guided behaviors such as the anterior ectosylvian visual cortex, striatum, and the deep layers of the superior colliculus, despite the fact that these structures themselves do not receive substantial direct projections from the visual cortical areas that are associated with the medial bank. Examination of the laminar location of the cells-of-origin of striate and extrastriate projections to LS using retrograde trace, WGA-HRP, revealed that the supragranular laminae of areas 17, 18 and 19 were the source of LS afferents whereas afferents from the other cortical areas (e.g., 20a, 20b, 21a, 21b, 7 and anterior ectosylvian visual area) were from both supra- and infragranular laminae. In addition, all LS subregions received intra-areal afferent projections from all LS cortical laminae. Thus, although rather clear hierarchical relationship between LS and visual cortical areas appears to exist, the interconnections among LS subregions provide no clear evidence of simple hierarchical relationships between regions LS or may have feed-forward and feed-back pathways.

Pretectofugal fibers from the nucleus of the optic tract in monkeys.

The nucleus of the optic tract (NOT) is the visuo-motor relay between the retina and preoculomotor structures in the pathway mediating optokinetic nystagmus (OKN). NOT lesions in monkeys produce no OKN toward the lesioned side. Then, efferent fibers from the NOT course through the brainstem and may reach the vestibular nucleus, which is proposed to be the final nucleus to the motor nucleus. In the present study, the tracer was injected through a micropipette in the NOT in four monkeys. Labeled terminals were observed ipsilaterally in the parabigeminal nucleus, superficial layers of the superior colliculus, dorsal and lateral terminal nuclei of the accessory optic system and pretectal nuclei and contralaterally in the NOT and superficial layers of the superior colliculus. Descending fibers from the NOT consisted of two major pathways: (1) fibers descended medially from the injection site through the reticularis pontis oralis to reach the lateral part of the ipsilateral nucleus reticularis tegmenti pontis; (2) fibers projecting into the dorsal cap of inferior olive, by far the greatest number of labeled fibers, descended ventrally along the lateral border of the reticularis pontis oralis and reached the medial lemniscus where they descended further and branched into the dorsolateral pontine nucleus, the lateral part of the nucleus reticularis tegmenti pontis, the peduncular pontine nucleus, the lateral pontine nucleus, the nucleus prepositus hypoglossi, the medial vestibular nucleus and finally the dorsal cap of the inferior olive. Consistent with the physiological data, the direct terminals to the medial vestibular nucleus could serve to drive the storage mechanisms and to produce OKN in the monkey.

Morphological analysis of the role of the neural tube and notochord in the development of somites.

The differentiation of avian somites and skeletal muscles, which themselves are derived from somites, was studied in ovo after the isolation of the unsegmented segmental plate from the notochord and/or neural tube by surgical operations at the level of the segmental plate. In each experiment, the newly formed somites had a normal histological structure, with an outer epithelial somite and core cells in the somitocoeles. Thereafter, the three derivatives of the somites (dermatome, myotome and sclerotome) reacted differently to the different operations. When the somites developed without the notochord, only somitocoele cells showed massive cell death, and muscles developed regardless of the presence or absence of the neural tube. On the contrary, no cell death was observed in any part of the somites that were formed with the neural tube or the notochord present, and muscle cells developed. However, in those embryos that retained only the notochord, striated muscles developed only in the lateral body wall. In each of the experimental operations, the surface ectoderm always covered the somites, and, regardless of the state of sclerotome and/or myotome differentiation, the dermatome always survived. These histological observations indicate that the first step in somite formation is independent of axial structures. The results further suggest that the notochord may produce diffusible factors that are necessary for the survival and further development of sclerotomal cells, and that both the neural tube and notochord can support muscle differentiation. However, it is likely that each structure has a relationship to the development of epaxial muscles and hypaxial muscles respectively. Furthermore, an intimate relationship may also exist between the surface ectoderm and the development of the dermatome.