Functional organization of the ventral lateral geniculate complex of the tree shrew (Tupaia belangeri): II. Connections with the cortex,thalamus, and brainstem

Connections of the ventral lateral geniculate complex (GLv) in the tree shrew were traced by anterograde and retrograde transport of WGA-HRP. The results buttress earlier findings that GLv in this species is composed of two main divisions, lateral and medial, each of which differs in its connections with the brainstem and cerebral cortex. The connections of the lateral division (GLv) suggest that it participates in visuosensory functions: it receives input from the retina, striate cortex, pretectum, and retino-recipient layers of the superior colliculus. These connections help clarify the identification of the internal and external subdivisions of GLv inasmuch as projections from both the superior colliculus and pretectum terminate in the external subdivision and each, in turn, receives a projection from the internal subdivision. Connections of the medial division suggest that this part of the nucleus is involved with visuomotor functions. Thus, the medio-caudal subdivision projects to the pontine nuclei, the prerubral field and the central lateral nucleus. The medio-caudal subdivision also receives projections from the lateral cerebellar nucleus, so that the GLv-ponto-cerebello-GLv loop involves mainly one subdivision of GLv. The medio-rostral subdivision receives projections from the pretectum and parietal cortex. Its output is directed primarily at the intermediate and deep layers of the superior colliculus. All of these targets of GLv, the pons, prerubral field, and deep layers of the superior colliculus, are known to play a role in the coordination of head and eye movements. Additional connections of GLv with the vestibular nuclei, intralaminar nuclei, hypothalamus, and facial motor nucleus are also described. © 1993 Wiley-Liss, Inc.     

Subcortical projections of six visual cortical areas in the owl monkey, Aotus trivirgatus.

Subcortical projections of six visual cortical areas (Areas 17 and 18, the Middle Temporal, Dorsomedial and Medial Areas, and the Posterior Parietal Region) in the owl monkey, Aotus trivigatus, were investigated with autoradiographic methods following injections of tritiated proline. No contralateral projections were demonstrated. While some brainstem structures received input from all six subdivisions of cortex, each cortical area appeared to exhibit its own unique pattern of subcortical projections. All six cortical areas were found to project to the superior and inferior divisions of the pulvinar, reticular nucleus of the thalamus, pretectum and superior colliculus. Other subcortical targets of one or more visual cortical areas were the basal ganglia, claustrum, zona incerta, one or more of the intralaminar nuclei, lateral posterior nucleus, pregeniculate nucleus, dorsal lateral geniculate nucleus, and pontine nuclei. Furthermore, details of corticofugal projections to the dorsal lateral geniculate nucleus, pretectum and superior colliculus varied with the cortical area studied. The projections to the reticular nucleus, pregeniculate nucleus, dorsal lateral geniculate nucleus, the inferior and a portion of the superior division of the pulvinar and the superior colliculus were found to be topographically organized. The targets of the subcortical projections were compared with those of the retina, as revealed by autoradiographic methods following tritiated proline injections of the eye and were found to overlap to varying extents in the superior colliculus, pretectum and dorsal lateral geniculate nucleus and to be segregated in the pregeniculate nucleus. The results substantiate the validity of previous studies in the owl monkey that suggest that the visual cortex is subdivided into several functionally distinct areas; and illustrate the complexity of corticofugal influence on visual processing.     

Thalamic connections of the ground squirrel superior colliculus and their topographic relations

The connections of the superior colliculus (SC) of the ground squirrel Spermophilus tridecemlineatus were studied with the horseradish peroxidase (HRP) method. Multiple pressure injections of HRP served to define the total pattern of SC projections while iontophoretic injections allowed differentiation of connections of the deep and superficial layers and determination of topographic relations of SC with its associated nuclei. The deep laminae were mainly connected with auditory, somatosensory and reticular regions of the brain, including the inferior colliculus, zona incerta, substantia nigra, mesencephalic central grey, pontine nuclei, spinal trigeminal nucleus, nucleus of the posterior commissure, thalamic reticular nucleus, raphe nuclei, lateral vestibular nucleus, the lateral superficial reticular formation of the medulla, the mesencephalic reticular formation, nucleus gracilis and the cervical spinal cord. The superficial laminae were connected with visual system structures. They were reciprocally connected with the dorsal and ventral lateral geniculate nuclei, the pretectum, nucleus lateralis posterior (LP), the parabigeminal nucleus and the contralateral SC. Connections between the SC and the dorsal lateral geniculate were topologic. LP was found to consist of three divisions: rostrolateral, rostromedial and caudal. SC was interconnected with the rostrolateral and caudal divisions. The connections between the SC and the rostrolateral division were topologic; those with the caudal division were not. The connections of the deep collicular layers in ground squirrels were similar to those which have been reported for cats and monkeys. The connections of the superficial laminae were more extensive than has been reported in other species. These elaborate interconnections indicate extensive interaction between primary retinal projection nuclei in the processing of visual information.     

An autoradiographic study of the efferent connections of the superior colliculus in the cat

The differential projections of the three main cellular strata of the superior colliculus have been examined in the cat by the autoradiographic method. The stratum griseum superficiale projects caudally to the parabigeminal nucleus and rostrally to several known visual centers: the nucleus of the optic tract and the olivary pretectal nucleus in the pretectum; the deepest C laminae of the dorsal lateral geniculate nucleus; the large-celled part of the ventral lateral geniculate nucleus; the posteromedial, large-celled part of the lateral posterior nucleus of the thalamus. Several of these projections are topographically organized. The stratum griseum profundum gives rise to most of the descending projections of the superior colliculus. Ipsilateral projections pass to both the dorsolateral and lateral divisions of the pontine nuclei, the cuneiform nucleus, and the raphe nuclei, and to extensive parts of the brainstem reticular formation: the tegmental reticular nucleus, and the paralemniscal, lateral, magnocellular, and gigantocellular tegmental fields. Contralateral projections descending in the predorsal bundle pass to the medial parts of the tegmental reticular nucleus and of some of the tegmental fields, the dorsal part of the medial accessory nucleus of the inferior olivary complex, and to the ventral horn of the cervical spinal cord. Ascending projections of the stratum griseum profundum terminate in several nuclei of the pretectum, the magnocellular nucleus of the medial geniculate complex and several intralaminar nuclei of the thalamus, and in the fields of Forel and zona incerta in the subthalamus. The strata grisea profundum and intermediale each have projections to homotopic areas of the contralateral superior colliculus, to the pretectum, and to the central lateral and suprageniculate nuclei of the thalamus. However, the stratum griseum intermediale has few or no descending projections.     

Subcortical projections to the centromedian and parafascicular thalamic nuclei in the cat

The primary objective of this study is to identify the totality of input to the centromedian and parafascicular (CM-Pf) thalamic nuclear complex. The subcortical projections upon the CM-Pf complex were studied in the cat with three different retrograde tracers. The tracers used were unconjugated horseradish peroxidase (HRP), horseradish peroxidase conjugated to wheat germ agglutinin (WGA-HRP), and rhodamine-labeled fluorescent latex microspheres (RFM). Numerous subcortical structures or substructures contained labeled neurons with all three tracing techniques. These labeled structures included the central nucleus of the amygdala; the entopeduncular nucleus; the globus pallidus; the reticular and ventral lateral geniculate nuclei of the thalamus; parts of the hypothalamus including the dorsal, lateral, and posterior hypothalamic areas and the ventromedial and parvicellular nuclei; the zona incerta and fields of Forel; parts of the substantia nigra including the pars reticularis and pars lateralis, and the retrorubral area; the pretectum; the intermediate and deep layers of the superior colliculus; the periaqueductal gray; the dorsal nucleus of the raphe; portions of the reticular formation, including the mesencephalic, pontis oralis, pontis caudalis, gigantocellularis, ventralis, and lateralis reticular nuclei; the nucleus cuneiformis; the marginal nucleus of the brachium conjunctivum; the locus coeruleus; portions of the trigeminal complex, including the principal sensory and spinal nuclei; portions of the vestibular complex, including the lateral division of the superior nucleus and the medial nucleus; deep cerebellar nuclei, including the medial and lateral cerebellar nuclei; and lamina VII of the cervical spinal cord. Moreover, the WGA-HRP and rhodamine methods (known to be more sensitive than the HRP method) revealed several afferent sources not shown by HRP: the anterior hypothalamic area, ventral tegmental area, lateral division of the superior vestibular nucleus, nucleus interpositus, and the nucleus praepositus hypoglossi. Also, the rhodamine method revealed labeled neurons in laminae V and VI of the cervical spinal cord.     

Functional organization of mustached bat inferior colliculus: II. Connections of the FM2 region

Echolocating bats estimate target distance by analyzing the time delay between frequency-modulated portions of their emitted ultrasonic vocalizations and the resultant echoes. In the companion paper we investigated, in the central nucleus of the inferior colliculus, the representation of the predominant second-harmonic frequency-modulated component (FM2) of the mustached bat biosonar signal (O'Neill et al.: J. Comp. Neurol. 283:000-000,'89). In the present paper we report the connections of this part of the colliculus, as determined by focal, iontophoretic injections of HRP following single-unit mapping of the FM2 representation. It was found that the major inputs to the FM2 region of the inferior colliculus come from the contralateral cochlear nucleus; ipsilaterally from the medial superior olive, periolivary nuclei, and ventral and intermediate nuclei of the lateral lemniscus; and bilaterally from the lateral superior olive and dorsal nucleus of the lateral lemniscus. This study identifies for the first time those specific regions of brainstem nuclei providing input to the central nucleus of the inferior colliculus that process FM2 information in the mustached bat. The primary outputs of the FM2 region project to the medial and dorsal divisions of the medial geniculate body. In sharp contrast to other mammals, we found little evidence of connections to the ventral division of the medial geniculate. Other regions receiving significant inputs from the FM2 area include the deep superior colliculus ipsilaterally and the ipsilateral lateral pontine nuclei. Some fibers also terminated near the midline in the dorsal midbrain periaqueductal gray. Sparse intrinsic connections were also seen to the ipsilateral dorsoposterior division of the central nucleus and to the contralateral inferior colliculus at a location homologous to the injection site in the anterolateral division. The finding that FM2 projections to the medial geniculate heavily favor the medial and dorsal divisions is consistent with the location of "FM-FM" delay-dependent facilitation neurons found by Olsen (Processing of Biosonar Information by the Medical Geniculate Body of the Mustached Bat, Pteronotus parnellii. Dissertation, Washington Univ., St. Louis, '86) in these divisions, and with thalamocortical projection patterns in this species. These findings demonstrate that for the FM portions of the biosonar signal, a transformation from a tonotopic form of processing to a more specialized, convergent pattern of organization occurs at the level of the inferior colliculus outputs.     

Direct synaptic connections of axons from superior colliculus with identified thalamo-amygdaloid projection neurons in the rat: Possible substrates of a subcortical visual pathway to the amygdala

The aim of the present study was to identify synaptic contacts from axons originating in the superior colliculus with thalamic neurons projecting to the lateral nucleus of the amygdala. Axons from the superior colliculus were traced with the anterograde tracers Phaseolus vulgaris leucoagglutinin or the biotinylated and fluorescent dextran amine “Miniruby.” Thalamo-amygdaloid projection neurons were identified with the retrograde tracer Fluoro-Gold. Injections of Fluoro-Gold into the lateral nucleus of the amygdala labeled neurons in nuclei of the posterior thalamus which surround the medial geniculate body, viz. the suprageniculate nucleus, the medial division of the medial geniculate body, the posterior intralaminar nucleus, and the peripeduncular nucleus. Anterogradely labeled axons from the superior colliculus terminated in the same regions of the thalamus. Tecto-thalamic axons originating from superficial collicular layers were found predominantly in the suprageniculate nucleus, whereas axons from deep collicular layers were detected in equal density in all thalamic nuclei surrounding the medial geniculate body. Double-labeling experiments revealed an overlap of projection areas in the above-mentioned thalamic nuclei. Electron microscopy of areas of overlap confirmed synaptic contacts of anterogradely labeled presynaptic profiles originating in the superficial layers of the superior colliculus with retrogradely labeled postsynaptic profiles of thalamo-amygdaloid projection neurons. These connections may represent a subcortical pathway for visual information transfer to the amygdala. J. Comp. Neurol. 403:158–170, 1999. © 1999 Wiley-Liss, Inc.     

Direct projections from the cerebellar nuclei to the superior colliculus in the rabbit: an HRP study

Cerebellar projections to the superior colliculus in the rabbit were studied by the anterograde and retrograde HRP methods. Cerebellotectal fibers arise mainly from the anterior and posterior interpositus nuclei and terminate contralaterally in layer VII, layer VI, layer V, and the deep tier of layer IV of the superior colliculus. Cerebellotectal fibers from the posterior interpositus nucleus originate from the lateral part of the nucleus and end chiefly in the caudal part of the superior colliculus. Cerebellotectal fibers from the anterior interpositus nucleus arise from the ventral part of the nucleus and terminate mainly in the rostromedial part of the superior colliculus. Some neurons in the lateral cerebellar nucleus also send fibers contralaterally to the intermediate and deep layers of the superior colliculus, especially to its rostral and lateral parts. Few, if any, cerebellotectal fibers arise from the medial cerebellar nucleus.     

Connections of the caudal cerebellar interpositus complex in a new world monkey (Cebus apella)

The afferent and efferent connections of the cerebellar interpositus complex were studied in a capuchin monkey (Cebus apella) that had received a transcannular horseradish peroxidase implant into the caudal portion of the anterior interpositus nucleus and posterior interpositus nucleus. While the heaviest anterogradely labeled ascending projections were observed to the contralateral ventral posterolateral nucleus of the thalamus, pars oralis (VPLo), efferent projections were also observed to the contralateral ventrolateral thalamic nucleus (VLc) and central lateral (CL) nucleus of the thalamic intralaminar complex, magnocellular (and to a lesser extent parvicellular) red nucleus, nucleus of Darkschewitsch, zona incerta, nucleus of the posterior commissure, lateral intermediate layer and deep layer of the superior colliculus, dorsolateral periaqueductal gray, contralateral nucleus reticularis tegmenti pontis and basilar pontine nuclei (especially dorsal and peduncular), and dorsal (DAO) and medial (MAO) accessory olivary nuclei, ipsilateral lateral (external) cuneate nucleus (LCN) and lateral reticular nucleus (LRN), and to a lesser extent the caudal medial vestibular nucleus (MVN) and caudal nucleus prepositus hypoglossi (NPH), and dorsal medullary raphe. The heaviest retrograde labeling was corticonuclear Purkinje cells in the paramedian cerebellar cortex lateral to the vermis of lobules IV-VIII. Otherwise, retrogradely labeled sources of afferents were predominantly contralateral in the dorsal, dorsomedial, paramedian, and peduncular sectors of the basilar pons, NRTP, and dorsal accessory (DAO) and medial accessory (MAO) of olivary nuclei, but were predominantly ipsilateral in the LCN, LRN, and in the medullary reticular formation along the roots of the hypoglossal (XII) cranial nerve. It appeared that the connections with the contralateral dorsal basilar pons, NRTP, DAO and MAO, and ipsilateral LCN and LRN are reciprocal.(ABSTRACT TRUNCATED AT 250 WORDS)     

Parabigeminal projections to the superior colliculus in the cat

Small amounts of horseradish peroxidase were injected into the superior colliculus of the cat and the distribution of labeled neurons in the parabigeminal nuclei was mapped. After injections placed dorsal to the stratum opticum in the superior colliculus, the parabigeminal nucleus is the only mesencephalic and/or rhombencephalic structure in which labeled neurons are observed. The number of labeled neurons in the parabigeminal nucleus increases after injections that include both the superficial and the deep layers of the superior colliculus. Each part of the superior colliculus receives projections from wide areas of both parabigeminal nuclei, although it also receives more abundant projections from one or more restricted parts of these nuclei. The anterior third of the parabigeminal nuclei is the part which sends the fewest projections to the superior colliculus. These projections terminate principally in the central and intermediate part of the contralateral colliculus, while a smaller number of fibers terminate in the lateral and rostral part of the ipsilateral colliculus. The intermediate third of the parabigeminal nuclei sends projections to all parts of the ipsilateral colliculus, but the greatest number of these goes to the contralateral colliculus. These contralateral projections terminate principally in the lateral parts of the contralateral colliculus, and in lesser number in its central and rostral, and medial and rostral areas. The posterior third of the parabigeminal nucleus sends scant efferents to wide areas of the ipsilateral and contralateral colliculi, and a dense projection to the medial and intermediate, medial and caudal and central and intermediate parts of the ipsilateral colliculus. There are also consistent projections from the posterior third of the parabigeminal nucleus to the central and rostral and medial and rostral parts of this ipsilateral colliculus. These results demonstrate a topographical organization of the parabigemino-tectal projections in the cat, as a pathway that facilitates the integration in the colliculus of visual impulses of different origin in the retina. This organization permits the modulation of the superior colliculus in its participation in both the extrageniculate visual system and in the regulation of eye and head orientation movements through the parabigeminal projections to the superficial and deep layers of the colliculus, respectively.     

Afferent connections of the thalamic intralaminar nuclei in the cat

Afferents to the central lateral (CL), paracentral (PC) and central medial (CE) intralaminar nuclei (ILN) from cortical and subcortical sites were studied in the cat. We utilized stereotaxically guided injections of HRP into the CL and PC nuclei and tritiated leucine injections into various visual, parietal and limbic areas of cortex to demonstrate these connections. In studying the relatively weak visual cortical projections to the ILN, we demonstrated projections from areas 19, 20a, 21a, 21b, AMLS, PMLS and PLLS. However, our HRP injections into the ILN often revealed only a few labeled cells in any of the above areas; therefore conclusions regarding the absence of projections to ILN from remaining visual cortical areas should be made cautiously. The ILN receive heavier projections from the frontal eye fields, cingulate cortex, splenial cortex, insular cortex, somatosensory areas SI and SII, auditory areas SF, AII, and Ep, and parietal areas 5 and 7. The most robust projections appear to be from from frontal eye fields, cingulate and parietal areas. No topography was apparent in the projections to the ILN. All cortical projections originate ipsilaterally from layers V and VI. Heavy subcortical projections to the ILN originate in the pretectum, superior colliculus, reticular formation, and periaqueductal grey. Fewer afferents arise from several other brainstem and thalamic nuclei.     

Subcortical projections to lateral geniculate and thalamic reticular nuclei in the hooded rat

Restricted injections either of horseradish peroxidase conjugated with wheat germ agglutinin, or of unconjugated horseradish peroxidase were made into hooded rats in order to distinguish subcortical sources of afferents to dorsal lateral geniculate nucleus from those to the adjacent visually responsive thalamic reticular nucleus, which modulates geniculate activity. Five “nonvisual” brainstem regions project to the dorsal lateral geniculate nucleus: mesencephalic reticular formation, dorsal raphe nucleus, periaqueductal gray matter, dorsal tegmental nucleus, and locus coeruleus. Projections are generally bilateral, but ipsilateral projections dominate. Of these regions, three also project ipsilaterally to the thalamic reticular nucleus: mesencephalic reticular formation, periaqueductal gray matter, and dorsal tegmental nucleus. Similar discrete injections of horseradish peroxidase into ventral lateral geniculate nucleus allowed a comparison of afferents to dorsal and ventral lateral geniculate nuclei. In addition to the five nonvisual brainstem regions which project to the dorsal division, the ventral lateral geniculate nucleus receives afferents from the perirubral reticular formation and the central gray matter at the thalamic level. The dorsal and ventral lateral geniculate nuclei receive substantially different afferents from subcortical visual centres. The dorsal division receives projections from superior colliculus, pretectum, and parabigeminal nucleus whereas the ventral division receives afferents from superior colliculus, additional pretectal nuclei, lateral terminal nucleus of the accessory optic system, and the contralateral ventral lateral geniculate nucleus.     

Differential projection patterns of superior and inferior collicular neurons onto posterior paralaminar nuclei of the thalamus surrounding the medial geniculate body in the rat

The thalamic nuclei at the medial border of the medial geniculate body (i.e. the suprageniculate nucleus, the medial division of the medial geniculate nucleus, the posterior intralaminar nucleus and the peripeduncular nucleus) which relay sensory information to the amygdala are thought to receive convergent input from multiple sites. In order to delineate the organization of these multimodal thalamic nuclei, the locations of superior and inferior collicular neurons projecting to these nuclei were studied by means of retrograde transport methods. Small injections of the tracer Miniruby were made into single paralaminar thalamic nuclei. Injections of Miniruby into the suprageniculate nucleus labelled predominantly neurons in the stratum opticum of the superior colliculus, whereas injections into the medial division of the medial geniculate body, the posterior intralaminar nucleus and the peripeduncular nucleus labelled predominantly neurons in the deep layers of the superior colliculus. These injections also labelled neurons in the inferior colliculus. The majority of retrogradely labelled neurons were found in the external nucleus of the inferior colliculus and here predominantly in layer 2. Injections focused onto the medial division of the medial geniculate body additionally labelled magnocellular neurons in layer 3 of the external nucleus and a few neurons in the central nucleus. More ventrally located injections, focused onto the posterior intralaminar and peripeduncular nucleus, almost exclusively labelled neurons in layer 1 of the external nucleus and the dorsal part of the dorsal nucleus. After injections into the suprageniculate nucleus, only neurons in layer 2 were found. Neurons in the central nucleus of the inferior colliculus were only found after injections that involved the medial division of the medial geniculate body. The present results suggest that, despite a considerable degree of convergence in this thalamic region, each of these thalamic nuclei receives a unique pattern of projections from the superior and inferior colliculi. It appears that the thalamic nuclei may be concerned mainly, but not exclusively, with a single sensory modality, and give rise to parallel multimodal and unimodal pathways to the amygdala.     

Topographic organization of the projections from cortical areas 17, 18, and 19 onto the thalamus,pretectum and superior colliculus in the cat

The distribution of cortical projections from areas 17, 18, and 19 to the lateral thalamus, pretectum, and superior colliculus was investigated with the autoradiographic tracing method. Cortical areas 17, 18 and 19 were demonstrated to project retinotopically and in register upon the dorsal lateral geniculate nucleus, medial interlaminar nucleus, lateral zone of the lateral posterior complex, nucleus of the optic tract and superior colliculus. Area 19 was shown to project retinotopically upon the pulvinar nucleus. Clear retinotopic organization was not demonstrable in the projections of areas 17, 18 and 19 to the reticular complex of the thalamus and ventral lateral geniculate nucleus, or in the projection of area 19 to the anterior pretectal nucleus. The cortical projections were employed to define the retinotopic organization of the nucleus of the optic tract, pulvinar nucleus, and lateral zone of the lateral posterior complex. The cortical projections show the vertical meridian to be represented caudally, with the lower visual field represented laterally, and the upper visual field medially, within the nucleus of the optic tract. The projections of area 19 to the pulvinar nucleus demonstrate the lower visual field to be represented rostrally and the upper visual field caudally in this nucleus; the vertical meridian to be represented at the lateral border and the visual field periphery to be represented at the medial border of the pulvinar nucleus. Cortical projections to the lateral zone of the lateral posterior complex demonstrate the lower visual field to be represented rostrally and the upper visual field caudally; the vertical meridian to be represented at the medial limit and the visual field periphery at the lateral border of the termination zones. On the basis of the experimental findings, a new terminology is introduced for the feline lateral posterior complex. Divisions are proposed which correspond to zones with demonstrably distinct afferent input. The pulvinar nucleus is defined by the distribution of projections from area 19. Three flanking divisions are defined within the lateral posterior complex; a lateral division recipient of projections from area 17, 18 and 19, an interjacent division recipient of projections of the superficial layers of the superior colliculus, and a medial division flanking the tectorecipient zone medially.     

Projections of the inferior colliculus in cat

Projections of the feline inferior colliculus were studied using the Nauta-Laidlaw method to demonstrate degenerating axons. A subtentorial stereotaxic approach was used to avoid corticofugal degeneration. Direct connections from the inferior colliculus to the anterior midline cerebellar cortex were observed. There is a topographical relationship of fibers of the brachium of the inferior colliculus and parabrachial region to the superior colliculus. A definite projection to the superior colliculi including a pathway through its commissure was found from the parabrachial region. A projection exists from the inferior colliculus to the dorsolateral portion of the central gray as far as the pretectum. There is a projection along the medial portion of the superior colliculus to the pretectum. Projections to the midbrain reticular formation, central gray and superior colliculi were substantial. Thalamic projections include a distribution of fibers to the magnocellular and rostral portion of the principal divisions of the medial geniculate body and to the lateral posterior thalamic nucleus. The rostral connections of the inferior colliculus with areas other than the medial geniculate body indicate that it may function in roles in addition to that of a mass somatomotor reflex center. Possible roles of the inferior colliculus in attention, habituation, and integration of corticovisual and auditory impulses are suggested.     

The projections of cells in different layers of the cat's visual cortex.

The projection of cells in different layers of several cortical visual areas in the cat were studied using the method of retrograde transport of horseradish peroxidase. Injections of the enzyme were made through a recording micropipette, making it possible to localize the injection site by physiological criteria. We found that layer VI cells projected to the alteral geniculate nucleus, while a distinct population of cells in layer V projected to the superior colliculus. Cells in layers II and III were tha major sources of ipsilateral cortico-cortical connections. This pattern of projection was consistent from one visual area to another. Pyramidal cells appeared to be the source of cortico-geniculate, cortico-collicular and cortico-cortical projections. The proportion of cells within a layer that terminated in a given site varied from layer to layer: apparently all of the large pyramids in layer V had terminals in the superior colliculus, about half of the pyramids in layer VI had terminals in the lateral geniculate nucleus, while only a small proportion of the pyramids in layers II and III had terminals in any single cortical area. The results indicated a remarkable specificity in the projections of the cortical layers. The cortical connections of the different cell types in layers A and A1 of the lateral geniculate nucleus were also examined: the cells that projected to area 17 were much more numerous and were on the average smaller than those that projected to area 18. Projections to the cortex were also found from the pulvinar, the medial interlaminar nucleus and the posterior nucleus. Direct connections were observed to the lateral geniculate nucleus from several midbrain reticular nuclei. Finally, projections were found to the superior colliculus from the zona incerta, the reticular nucleus of the thalamus and the ventral lateral geniculate nucleus.     

Anterograde degeneration study of the superior colliculus in Tupaia glis: evidence for a subdivision between superficial and deep layers

The large, well developed superior colliculus of the tree shrew with its highly differentiated layers is ideal for analyzing the connections of individual layers. Our most significant finding concerns the differences between the projections of the superficial and deep layers. Lesions limited to those strata which receive projections from the retina and striate cortex (superficial 3 layers) result in terminal degeneration almost exclusively within the pulvinar, the pretectal area, and the dorsal and ventral lateral geniculate nuclei. In each case, the greatest amount of degeneration was present in the pulvinar, supporting previous suggestions that the tecto-pulvinar pathway conveys visual information. In sharp contrast, lesions limited to the deep layers which receive multimodal input from nonstriate areas of the neocortex and from a variety of subcortical centers, produce terminal degeneration in entirely different thalamic nuclei — certain intralaminar nuclei, the subthalamic region, and a region homologous to the posterior nuclear group of Rose and Woolsey ('49). The deep lesions also result in terminal degeneration within the inferior colliculus, the parabigeminal nucleus, the reticulo-tegmental nucleus, and the inferior olivary nucleus, as well as in the brainstem reticular formation. Finally, deep lesions produced scattered degenerating fibers in the motor facial nucleus. Our results favor a division of the tree shrew superior colliculus into superficial and deep portions based on strikingly different projection patterns and may be useful in resolving certain problems of thalamic homology.     

Retinal projections in the house musk shrew, Suncus murinus, as determined by anterograde transport of WGA-HRP.

Retinal projections in the house musk shrew (Suncus murinus) were determined by the anterograde transport of wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP). Unilateral injection of WGA-HRP into the vitreous body resulted in the terminal labeling of the optic projections in the suprachiasmatic nucleus (SCH), the ventral (CGLv) and dorsal (CGLd) lateral geniculate nuclei, the intergeniculate leaflet (IGL), the pretectum, the superficial layers of the superior colliculus (CS), and the dorsal terminal nucleus (DTN) of the accessory optic system (AOS). Labeling of the SCH was bilateral, with ipsilateral predominance, and covered the whole dorsoventral extent of the nucleus. Immunohistochemical studies revealed that VIP-like immunoreactive neurons and fibers were present in almost all parts of the SCH. No hypothalamic regions other than the SCH received the optic fibers. The ipsilateral projections to the CGLv, CGLd, and IGL were sparse, and a considerable number of uncrossed retinal fibers were found in the pretectal olivary nucleus. No retinal projections to the lateral posterior thalamic nucleus (LP) were found. Ipsilateral optic fibers projected sparsely to the medial part of the CS. The AOS consisted of a small DTN with a very few crossed retinal projections but no lateral and medial terminal nuclei. In addition, the AOS had no inferior fascicle.     

Connections of the pretectum in the cat.

The connections of the pretectal complex in the cat have been examined by anatomical methods which utilize the anterograde axonal transport of tritiated proteins or the retrograde axonal transport of the enzyme horseradish peroxidase. Following injections of tritiated amino acids into the eye, label can be seen in the contralateral and ipsilateral nucleus of the optic tract and olivary nucleus where it appears as two or three finger-like strips. Following large injections of tritiated amino acids into the pretectal complex transported label accumulates ipsilaterally in a region dorsolateral to the red nucleus, the central and pericentral divisions of the tegmental reticular nucleus, the intermediate layers of the superior colliculus, the nucleus of Darkschewitch, the thalamic reticular nucleus, zona incerta and fields of Forel, the central lateral nucleus, the pulvinar nucleus and the ventral lateral geniculate nucleus. Contralaterally label accumulates in the nucleus of the posterior commissure, the interstitial nucleus of Cajal, the anterior, posterior and medial pretectal nuclei, and the ventral lateral geniculate nucleus From smaller injections, more or less well confined to single nuclei, the following patterns of connections are demonstrated. The nucleus of the optic tract projects to the ipsilateral ventral lateral geniculate nucleus and pulvinar nucleus and to the contralateral nucleus of the posterior commissure. The anterior pretectal nucleus projects to the ipsilateral central lateral nucleus, the reticular nucleus, zona incerta, fields of Forel, the region dorsolateral to the red nucleus and to the contralateral anterior pretectal nucleus. The posterior pretectal nucleus seems to project only to the ipsilateral reticular nucleus and zona incerta. The central tegmental fields deep to the pretectum project to the tegmental reticular nucleus of the brainstem. When the injection involves the nucleus of the posterior commissure label is seen in the ipsilateral nucleus of Darkschewitch, and in the contralateral nucleus of the posterior commissure and interstitial nucleus of Cajal but no nucleus of the pretectum could be positively identified as projecting to any of the motor nuclei of cranial nerves III, IV, and VI. Following large injections of horseradish peroxidase into the pretectal complex, labeled cells are seen in the superficial layers of the ipsilateral superior colliculus, in the ipsilateral ventral lateral geniculate nucleus, reticular nucleus and zona incerta and in the contralateral anterior, medial and posterior pretectal nuclei, nucleus of the optic tract and ventral lateral geniculate nucleus.     

Efferent projections of the intergeniculate leaflet and the ventral lateral geniculate nucleus in the rat

The intergeniculate leaflet (IGL) and the ventral lateral geniculate nucleus (VLG) are ventral thalamic derivatives within the lateral geniculate complex. In this study, IGL and VLG efferent projections were compared by using anterograde transport of Phaseolus vulgaris-leucoagglutinin and retrograde transport of FluoroGold. Projections from the IGL and VLG leave the geniculate in four pathways. A dorsal pathway innervates the thalamic lateral dorsal nucleus (VLG), the reuniens and rhomboid nuclei (VLG and IGL), and the paraventricular nucleus (IGL). A ventral pathway runs through the geniculohypothalamic tract to the suprachiasmatic nucleus and the anterior hypothalamus (IGL). A medial pathway innervates the zona incerta and dorsal hypothalamus (VLG and IGL); the lateral hypothalamus and perifornical area (VLG); and the retrochiasmatic area (RCA), dorsomedial hypothalamic nucleus, and subparaventricular zone (IGL). A caudal pathway projects medially to the posterior hypothalamic area and periaqueductal gray and caudally along the brachium of the superior colliculus to the medial pretectal area and the nucleus of the optic tract (IGL and VLG). Caudal IGL axons also terminate in the olivary pretectal nucleus, the superficial gray of the superior colliculus, and the lateral and dorsal terminal nuclei of the accessory optic system. Caudal VLG projections innervate the lateral posterior nucleus, the anterior pretectal nucleus, the intermediate and deep gray of the superior colliculus, the dorsal terminal nucleus, the midbrain lateral tegmental field, the interpeduncular nucleus, the ventral pontine reticular formation, the medial and lateral pontine gray, the parabrachial region, and the accessory inferior olive. This pattern of IGL and VLG projections is consistent with our understanding of the distinct functions of each of these ventral thalamic derivatives.