Optic tectum of the eastern garter snake, Thamnophis sirtalis. I. Efferent pathways

Extracellular, iontophoretic injections of horseradish peroxidase were used to anterogradely fill axons efferent from the optic tectum in garter snakes. The tectal efferent pathways consist of six axon types with distinct projections and terminal morphologies. Tectogeniculate axons pass into the diencephalon via the optic tract, bearing collaterals that form spatially restricted, rodlike arbors in the pretectum, the ventral lateral geniculate nucleus, and the ventrolateral nucleus. Tectoisthmi axons exit the tectum as a thin-caliber component of the ventral tectobulbar tract. They form spatially restricted, spherical arbors within nucleus isthmi. Tectoisthmobulbar axons also give rise to small, spherical arbors within nucleus isthmi, but the parent axons continue caudally into the pontine and medullary reticular formation issuing many short collateral branches. Tectorotundal axons reach the diencephalon via the tectothalamic tract and give rise to fine terminal collaterals in the nucleus of the tectothalamic tract ipsilaterally and in nucleus rotundus bilaterally. Single axons form sheetlike terminal fields that span the rostrocaudal extent of nucleus rotundus. Ipsilateral tectobulbar axons descend into the midbrain tegmentum where they issue several thick collaterals that terminate widely throughout the nucleus lateralis profundus mesencephali. The parent axon continues caudally giving off several widely spreading collaterals within the pontine and medullary reticular formation. Crossed tectobulbar axons enter the dorsal tectobulbar tract and cross the midline to form the predorsal bundle. Single axons give rise to terminal collaterals in the nucleus lateralis profundus mesencephali bilaterally, the contralateral pontine and medullary reticular formation, and the intermediate gray of the cervical spinal cord.     

Visual and electrosensory circuits of the diencephalon in mormyrids: an evolutionary perspective

Mormyrids are one of two groups of teleost fishes known to have evolved electroreception, and the concomitant neuroanatomical changes have confounded the interpretation of many of their brain areas in a comparative context, e.g., the diencephalon, where different sensory systems are processed and relayed. Recently, cerebellar and retinal connections of the diencephalon in mormyrids were reported. The present study reports on the telencephalic and tectal connections, specifically in Gnathonemus petersii, as these data are critical for an accurate interpretation of diencephalic nuclei in teleosts. Injections of horseradish peroxidase into the telencephalon retrogradely labeled neurons ipsilaterally in various thalamic, preglomerular, and tuberal nuclei, the nucleus of the locus coeruleus (also contralaterally), the superior raphe, and portions of the nucleus lateralis valvulae. Telencephalic injections anterogradely labeled the dorsal preglomerular and the dorsal tegmental nuclei bilaterally. Injections into the optic tectum retrogradely labeled neurons bilaterally in the central zone of area dorsalis telencephali and ipsilaterally in the torus longitudinalis, various thalamic, pretectal, and tegmental nuclei, some nuclei in the torus semicircularis, the nucleus of the locus coeruleus, the nucleus isthmi and the superior reticular formation, basal cells in the ipsilateral valvula cerebelli, and eurydendroid cells in the contralateral lobe C4 of the corpus cerebelli. Weaker contralateral projections were also observed to arise from the ventromedial thalamus and various pretectal and tegmental nuclei, and from the locus coeruleus and superior reticular formation. Tectal injections anterogradely labeled various pretectal nuclei bilaterally, as well as ipsilaterally the dorsal preglomerular and dorsal posterior thalamic nuclei, some nuclei in the torus semicircularis, the dorsal tegmental nucleus, nucleus isthmi, and, again bilaterally, the superior reticular formation. A comparison of retinal, cerebellar, tectal, and telencephalic connections in Gnathonemus with those in nonelectrosensory teleosts reveals several points: (1) the visual area of the diencephalon is highly reduced in Gnathonemus, (2) the interconnections between the preglomerular area and telencephalon in Gnathonemus are unusually well developed compared to those in other teleosts, and (3) two of the three corpopetal diencephalic nuclei are homologues of the central and dorsal periventricular pretectum in other teleosts. The third is a subdivision of the preglomerular area, rather than an accessory optic or pretectal nucleus, and is related to electroreception. The preglomerulo-cerebellar connections in Gnathonemus are therefore interpreted as uniquely derived characters for mormyrids.     

Connections of the tectum opticum in two urodeles, Salamandra salamandra and Bolitoglossa subpalmata, with special reference to the nucleus isthmi

Tectal connections were studied in two urodele species following horseradish peroxidase injections into the tectum opticum. In both species retrogradely labelled cells were observed: ipsilaterally in the corpus striatum, lateral amygdala, ventral and dorsal thalamus and nucleus of DARKSCHEWITSCH--bilaterally in the pretectal nucleus, dorsal tegmentum and nucleus reticularis medius--contralaterally in the tectum opticum and area octavo lateralis. Besides these nuclei the nucleus isthmi was bilaterally labelled. Rostral efferent projections of the tectum opticum terminated in the ipsilateral pretectal area and the ipsilateral dorsal and ventral thalamus ipsilaterally coursing to the contralateral tectum via the commissura postoptica. Caudal efferents formed the bilaterally organized tecto-bulbar tracts innervating the rhombencephalon. Comparison of the results of a series of tectal horseradish peroxidase injections differing in depth, tangential extension and location, indicated that tectal afferents from the telencephalon, the contralateral tectum opticum and the medulla were sparse and widely branching. Projections of the telencephalon and all diencephalic nuclei terminated deep in the rostral tectum opticum. Projections of the medulla terminated preferentially deep in the caudal tectum opticum. The tecto-isthmic projection was highly topographic forming a layered terminal field lateral to the nucleus isthmi. The isthmo-tectal projection innervated the whole tectum opticum on the ipsilateral side and was highly topographic. On the contralateral side the caudal part of the tectum opticum was not innervated. The isthmo-tectal fibers terminated superficially in the tectum opticum on both sides of the brain. The nucleus isthmi identified here is proposed to be homologe to that of other vertebrates.     

Afferents to the optic tectum of the leopard frog: an HRP study.

Following unilateral HRP injections in the optic tectum of Rana pipiens, HRP-positive cells were seen in three pretectal nuclei: bilaterally in the dorsal posterior nucleus; in the dorsal half of the ipsilateral posterior nucleus; and ipsilaterally in the large-called pretectal nucleus. HRP-positive cells were also seen ipsilaterally in the anterodorsal, posterodorsal and posteroventral tegmental fields, the nucleus isthmi, and the dorsal gray columns of the cervical spinal cord; bilaterally in the suprapeduncular nucleus, a paramedian cell group dorsal to the interpeduncular nucleus; and in the deep layers of the contralateral tectum. In addition, evidence for a bilateral ventral preopto-tectal projection was seen in half the experimental animals. No tectal afferents from telencephalic or rostal thalamic areas were seen. Both the ascending and descencing tectal efferent fibers were also filled with reaction product. The pale reaction indicative of terminating tectal efferents was seen in the dorsal pretectum, partially overlapping the lateral nucleus and uncinate neuropil; in the core of nucleus isthmi; and in the superior olive.     

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.     

Afferent and efferent connections of the optic tectum in the carp (Cyprinus carpio L.)

The afferent and efferent connections of the tectum opticum in the carp (Cyprinus carpio L.) were studied with the HRP method. Following iontophoretic peroxidase injections in several parts of the tectum anterograde transport of the enzyme revealed tectal projections to the lateral geniculate nucleus, dorsal tegmentum, pretectal nuclei, nucleus rotundus, torus longitudinalis, torus semicircularis, nucleus isthmi, contralateral tectum and to the mesencephalic and bulbar reticular formations.Tectal afferents were demonstrated by retrograde HRP transport in the area dorsalis pars centralis of the telencephalon, torus longitudinalis, torus semicircularis, nucleus isthmi, nucleus profundus mesencephali, several pretectal nuclei, dorsomedial and dorsolateral thalamic nuclei, nucleus of the posterior commissure, mesencephalic and bulbar reticular nuclei and nucleus ruber. Visuo-cerebellar circuitry was investigated by means of peroxidase injections in the various parts of the cerebellum. These experiments revealed indirect retino- and tecto-cerebellar pathways via the pretectal nuclei and the nucleus isthmi.     

Projections of the optic tectum in the longnose gar,lepisosteus osseus

Efferent projections of the optic tectum were studied with the anterograde degeneration method in the longnose gar. Ascending projections were found bilaterally to 3 pretectal nuclei — the superficial pretectal nucleus, nucleus pretectalis centralis and nucleus pretectalis profundus — and to a number of targets which lie further rostrally — the central posterior nucleus, dorsal posterior nucleus, accessory optic nucleus, nucleus ventralis lateralis, nucleus of the ventral optic tract, rostral part of the preglomerular complex, suprachiasmatic nucleus, anterior thalamic nucleus, nucleus ventralis medialis, nucleus intermedius, nucleus prethalamicus and rostral entopeduncular nucleus. Projections of the tectum reach the contralateral side via the supraoptic decussation and are less dense contralaterally than ipsilaterally. Descending projections resulting from tectal lesions include: (1) a tectal commissural pathway to the core of the torus longitudinalis bilaterally and the contralateral tectum and torus semicircularis; and (2) a pathway leaving the tectum laterally from which fibers terminate in the ipsilateral torus semicircularis, an area lateral to the nucleus of the medial longitudinal fasciculus, lateral tegmental nucleus, nucleus lateralis valvulae, nucleus isthmi and the reticular formation. A component of this bundle decussates at the level of the lateral tegmental nucleus to project to the contralateral reticular formation.

On the basis of comparisons of these findings with the pattern of retinal projections in gars and other data, it is argued that the nuclei previously called the lateral geniculate and rotundus in fish are not the homologues of the nuclei of those names in land vertebrates but are rather pretectal cell groups. The overall organization of both retinal and tectal projections in gars is strikingly similar to that in land vertebrates; at present, the best candidate for a rotundal homologue is the dorsal posterior nucleus.     

Tectal connections in Python reticulatus

The origins of the axons terminating in the mesencephalic tectum in Python reticulatus were examined by unilateral tectal injections of horseradish peroxidase. Kutrogradely labeled cells were observed bilaterally throughout the spinal cord in all subdivisions of the trigeminal system, with the exception of nucleus principalis, which showed labeled cells only on the ipsilateral side. Labeling of the reticular formation occurred bilaterally in nucleus reticular is interiormagnocellularis, nucleus reticularis lateralis, nucleus reticularis medius and the mesencephalic reticular formation. The tectum also receives bilateral projections from the dorsal tegmentaJ field, the nucleus of the lateral lemniscus and nucleus isthmi, and ipsilateral projections from nucleus profundus mesencephali. A few labeled cells were found ipsilaterally in the locus coeruleus and in nuclei vestibulares ventrolateralis and centromedialis. In the diencephalon labeled cells were observed ipsilaterally in nucleus ventrolateralis thalami, nucleus ventromedialis thalami, nucleus suprapeduncularis, and in the dorsal and ventral lateral geniculate nuclei. Bilateral labeling was observed in nucleus periventricularis hypo-thalami. Furthermore, labeling was ipsilaterally present in the ventral telen-cephalic areas. The tectum in Python reticulatus receives a wide variety of afferent connections which confirm the role of the tectum as an integration center of visual and exteroceptive information.     

Tectal projections of an infrared sensitive snake,Crotalus viridis

Crotaline snakes have detectors for infrared radiation and this information is projected to the optic tectum in a spatiotopic manner. The tectal projections were examined in Crotalus viridis with the use of silver methods for degenerating fibers and the autoradiographic and horseradish peroxidase tracing methods. Large lesions included all of the tectal layers but not the underlying structures. Projections to the thalamus include a sparse input to the ipsilateral ventral and dorsal lateral geniculate nuclei, the ventromedial nucleus, and nucleus lentiformis thalami. Nucleus rotundus was not detected. The projections to the pretectal nuclei are primarily ipsilateral to the nucleus lentiformis mesencehali and pretectal nucleus. At the level of the mesencephalon, tectal efferents are bilateral to nucleus profundus mesencephali and the tegmentum. There is minimal input to the contralateral deep tectal layers. There are ispilateral terminations in a nucleus identified as the posterolateral tegmental nucleus. Descending fibers include the two major tracts—the ventral tectobulbar tract that terminates in the ipsilateral lateral reticular formation and the predorsal bundle that distributes throughout the contralateral medial reticular formation. Two small descending tracts were noted—the intermediate and dorsal tectobulbar tracts. All of these descending tracts appear to terminate by the time they reach the caudal medulla. After superficial lesions terminals could be found in the ventral lateral geniculate nucleus, the nucleus profundus mesencephali, and the posterolateral tegmental nucleus; the two major descending tracts contained degenerated fibers as well. The areas receiving tectal input in Crotalus were compared to those of other reptiles and discussed.     

Tectal efferents in the banded water snake,Natrix sipedon

Visual information reaches the dorsal thalamus by two distinct routes in most reptiles. Retinal efferents terminate directly in the dorsal lateral geniculate nucleus (DLGN). Retinal information is also channeled indirectly through the tectum to nucleus rotundus. Retinal projections to DLGN and tectum are also well esablished in snakes, but the status of the tecto-rotundal link of the indirect visual pathway is uncertain. Thus, tectal efferents were studied with Fink-Heimer methods in banded water snakes (Natrix sipedon). The tectum gives rise to crossed and uncrossed projections to the brainstem reticular formation. Commissural connections are effected with the contralateral tectum via the tectal and osterior commissures. tectum projects densely to the ipsilateral basal optic nucleus. Bilateral ascending projections reach the pretectal area, nucleus lentiformis mesencephali, lateral habenular nuclei, and posterodorsal nuclei. Ascending projections reach the ventral lateral geniculate and suprapeduncular nuclei. there is a diffuse projection to the central part of the caudal thalamus and a dense, bilaternal projection to the DLGN. These results indicate that the relation of the tectum to the dorsal thalamus is different in snakes than in other reptiles. Nucleus rotundus is either absent or poorly differentiated and there is a strong convergence of the direct and indirect visual pathways at DLGN.     

Localization of neurons afferent to the optic tectum in longnose gars

Afferent pathways to the optic tectum in the longnose gar were determined by unilateral tectal injections of HRP. Retrogradely labeled cells were observed in the ipsilateral caudal portion of the rostral entopeduncular nucleus and bilaterally in the rostral half of the lateral zone of area dorsalis of the telencephalon. The following diencephalic cell groups were also labeled following tectal injections: the ipsilateral anterior, ventrolateral, and ventromedial thalamic nuclei, the periventricular pretectal nucleus, and the central pretectal nucleus (bilaterally); the ventromedial thalamic and central pretectal nuclei revealed the largest number of labeled cells. At midbrain levels, retrogradely labeled cells were seen in the ipsilateral torus longitudinalis, nucleus isthmi, and accessory optic nucleus; cells were labeled bilaterally in the torus semicircularis and a rostral tegmental nucleus. Only a few cells were labeled in the contralateral optic tectum, suggesting that few of the fibers of the intertectal commissure are actually commissural to the tectum. At hindbrain levels, retrogradely labeled cells were seen bilaterally in the locus coeruleus, the superior, medial, and inferior reticular formations, the eurydendroid cells of the cerebellum, and the nucleus of the descending trigeminal tract; the contralateral dorsal funicular nucleus also exhibited labeling. Clearly, the tectum in gars receives a substantial number of nonvisual afferents from all major brain areas, most of which have been reported in other vertebrates. The functional significance of these afferent sources and their probable homologues in other vertebrate groups are discussed.     

Tectoreticular pathways in the turtle, Pseudemys scripta. I. Morphology of tectoreticular axons

Tectoreticular projections in turtles were examined by reconstructing from serial sections axons that were anterogradely filled with horseradish peroxidase after tectal injections. Three tectoreticular pathways each contain extensively collateralized axons. The crossed dorsal pathway (TBd) contains large and small caliber axons. After leaving the tectum, TBd axons emit collaterals into the ipsilateral profundus mesencephali rostralis and then give off a main rostral branch that bears secondary collaterals in the ipsilateral interstitial nucleus of the medial longitudinal fasciculus and the suprapeduncular nucleus. The main trunks cross the midline and descend in the predorsal bundle, generating collaterals at regular intervals. These terminate mostly in the medial half of the reticular core from the midbrain to the caudal medulla. Axons in the uncrossed intermediate pathway also emit collaterals into a midbrain reticular nucleus (profundus mesencephali caudalis) and often have a thick rostral branch. The main caudal trunks, however, remain ipsilateral and travel in a diffuse, laterally placed tract, where each emits a long series of collaterals into the lateral half of the reticular core. The uncrossed ventral pathway (TBv) contains medium and small caliber axons. TBv axons often have collaterals within the tectum and apparently lack main rostral branches. Their caudal trunks run in the tegmental neuropile below the TBi where they collateralize less exuberantly than do TBd and TBi axons. The morphology of axons in all three pathways suggests that projections from disjunct tectal loci converge at many rostrocaudal levels within the reticular formation. This point was examined explicitly in experiments in which two disjunct injections were placed in one tectal lobe. Intermediate pathway axons traced from the two loci initially formed two distinct bundles but then intermingled in the reticular formation.     

Afferent connections of the optic tectum in lampreys: an experimental study

Tectal afferents were studied in adult lampreys of three species (Ichthyomyzon unicuspis, Lampetra fluviatilis, and Petromyzon marinus) following unilateral BDA injections into the optic tectum (OT). In the secondary prosencephalon, neurons projecting to the OT were observed in the pallium, the subhipoccampal lobe, the striatum, the preoptic area and the hypothalamus. Following tectal injections, backfilled diencephalic cells were found bilaterally in: prethalamic eminence, ventral geniculate nucleus, periventricular prethalamic nucleus, periventricular pretectal nucleus, precommissural nucleus, magnocellular and parvocellular nuclei of the posterior commissure and pretectal nucleus; and ipsilaterally in: nucleus of Bellonci, periventricular thalamic nucleus, nucleus of the tuberculum posterior, and the subpretectal tegmentum, as well as in the pineal organ. At midbrain levels, retrogradely labeled cells were seen in the ipsilateral torus semicircularis, the contralateral OT, and bilaterally in the mesencephalic reticular formation and inside the limits of the retinopetal nuclei. In the hindbrain, tectal projecting cells were also bilaterally labeled in the dorsal and lateral isthmic nuclei, the octavolateral area, the sensory nucleus of the descending trigeminal tract, the dorsal column nucleus and the reticular formation. The rostral spinal cord also exhibited a few labeled cells. These results demonstrate a complex pattern of connections in the lamprey OT, most of which have been reported in other vertebrates. Hence, the lamprey OT receives a large number of nonvisual afferents from all major brain areas, and so is involved in information processing from different somatic sensory modalities.     

Observations on the projections and intrinsic organization of the pigeon optic tectum: an autoradiographic study based on anterograde and retrograde, axonal and dendritic flow.

Three aspects of the labelling pattern seen after the injection of 13 different radioactive amino acids into the pigeon optic tectum have been described: The efferent projections of the optic tectum; the specific labelling of two pathways; and the dendritic organisation of tectal layer III neurons based on the retrograde and anterograde movement of label within these dendrites. Discrete injections of tritiated amino acid that involved all or only the superficial tectal layers suggested that layer III gave rise to the massive non-topographically organised and bilateral projections (fibers crossing within the decussato supraoptica ventrlis) upon the nuclei rotundus, subpraetectalis and interstitio-praetecto-subpraetectalis and to the ipsilaterally directed pathways terminating within the nuclei praetectalis, triangularis, subrotundus, dorsolateralis anterior thalami, posteroventralis and ventrolateralis thalami. Layer III neurons may also be the source of efferents to the posterior dorsolateral thalamus (the layer III pathway), the pontine grey and, bilaterally to the reticular formation and of the layer IV or tectal commisural pathway terminating within the contralateral tectal cortex. In contrast projections originating from layer II were generally topographically organised and terminated either within certain of the isthmic nuclei (n. isthmi pars parvocellularis, n. isthmo-opticus and n. semilunaris) or ran within layer I (layer I pathways) to end in the pretectum (griseum tectale) and ventral thalamus (n. ventrolateralis thalami, n. geniculatus, pars ventralis). A small projection from layer II upon the ipsilateral nucleus rotundus may also be present. Triated serine and tyrosine were found to be particularly effective in labeling perikarya as well as axons and terminals. The layer I pathway could be selectively labelled after tectal injections of 3H-GABA while the cell bodies of Ipc neurons were labelled in a retrograde fashion after tectal injections of 3H-glycine, serine or alanine. Intrinsic tectal labelling was found by correlation with Golgi material to reflect both anterograde and retrograde transport of label within dendrites of layer III cells. Anterograde movement of label indicated that the terminal portions of layer III cell dendrites ended in an orderly radial arrangement within sublayers IIb and IId, while the retrograde movement of label resulted in the labelin of layer III perikarya outside the injection field.     

The effects of ablation of visual cortex in neonatal rabbits on the organization of retinothalamic and retinopretectal projections

Primary visual cortex was ablated unilaterally in neonatal rabbits. Following a survival of 2-4 months, retrograde degeneration of the dorsal lateral geniculate nucleus (LGd) was assessed, and reorganization of retinofugal pathways was studied using methods of anretrograde transport of [3H]proline or of horseradish peroxidase. A complete lesion of primary visual cortex resulted in complete retrograde degeneration of the LGd with no sparing of any class of neurons. The terminations of retinofugal axons in the pretectum and thalamus were compared with those observed in normal animals. No major reorganization of ipsilateral retinofugal projections was observed in either the thalamus and pretectum ipsilateral to the ablated cortex, or in the thalamus and pretectum contralateral to the ablated cortex. However, contralateral retinofugal projections to the thalamus and to the pretectum ipsilateral to the ablated cortex were significantly different from normal. In the thalamus, the projections to the lateral posterior nucleus were expanded in area and increased in density. In the pretectum, the projections to the rostral pretectal areas were greatly increased in area, especially in the region of the olivary pretectal nucleus and posterior pretectal nucleus. However, the density of these projections was not increased relative to normal. Consideration of these results in relation to other published data on the anatomical consequences of neonatal visual cortex lesions, both in mammals which show behavioral sparing following neonatal visual cortex lesions and in mammals which, like the rabbit, show no behavioral sparing, suggests that: (1) behavioral sparing may correlate with patterns of survival or death of neurons in the thalamus and retina; and (2) reorganization of retinofugal pathways is not necessarily associated with behavioral sparing.     

Evidence of sub-collicular auditory projections to the medial geniculate nucleus in the cat: An autoradiographic and horseradish peroxidase study

Connections of a posteromedial region of the ventral nucleus of the lateral lemniscus were examined in the cat using the autoradiographic tracing method. This sub-collicular region previously had been shown, using retrograde transport of horseradish peroxidase, to send axons to the superior colliculus¹⁰. The autoradiographic findings revealed that many axons from the posteromedial region of the ventral nucleus of the lateral lemniscus that entered the superior colliculus continued into the midbrain reticular formation. Moreover, other axons traced rostral to the inferior colliculus into the thalamus ended in the medial geniculate nucleus, bilaterally. Experiments in which horseradish peroxidase was placed in the medial geniculate nucleus retrogradely labeled the large neurons in the posteromedial region supporting the autoradiographic observations. Other sub-collicular regions also contained labeled cells in these cases, including the main body of the ventral nucleus of the lateral lemnicus and scattered cell groups around the superior olivary complex.     

Somatomotor connectivity in the midbrain of Rana pipiens

The mesencephalic oculomotor nuclei of Rana pipiens and their surrounding cell groups were investigated using the anterograde and retrograde transport of horseradish peroxidase and Golgi techniques. The cell groups surrounding the oculomotor and trochlear nuclei were divided into the nucleus interstitialis (nInt) groups A, B, and C, the basal optic nucleus, and the nucleus reticularis tegmenti. Afferents to the ventral mesencephalon originate from the retina and from vestibular, cerebellar, visual, and accessory oculomotor nuclei. These afferents produce a sequence of terminal arborizations in which visual afferents are found in the outer neuropil, and accessory oculomotor, vestibular, and cerebellar afferents are found along the inner neuropil and central gray. The oculomotor neurons in anurans have extensive dendritic fields, extending to the outer margins of the neuropils, as do many large cells along the margin of nInt. Other neurons in nInt have dendritic fields restricted to the proximal portions of the neuropil. Efferents from nInt area A project to the cerebellum and bilaterally to the spinal cord. Area B nInt projects to the ipsilateral spinal cord, contralateral nInt, pretectal nucleus lentiformis mesencephali, and ipsilateral trochlear nucleus. Efferents from area C nInt reach the deep tectal layers and ipsilateral spinal cord. The outer portions of the neuropil contain the nucleus of the basal optic root which comprises ganglionic elongate and stellate neurons and projects to the pretectum. In the center of the neuropil peri-nBOR neurons have dendrites directed towards the visual terminal fields and axons towards the central gray and oculomotor neurons. The nucleus reticularis tegmenti receives afferents from the tectum and lateral forebrain bundle and projects to the deep tectal layers. In anurans, the oculomotor neurons receive a variety of visual, somatic, and vestibular afferents and appear relatively undifferentiated, whereas the nInt appears more developed.     

Putative cholinergic projections from the nucleus isthmi and the nucleus reticularis mesencephali to the optic tectum in the goldfish (Carassius auratus)

The nucleus isthmi of fish and amphibians has reciprocal connections with the optic tectum, and biochemical studies suggested that it may provide a major cholinergic input to the tectum. In goldfish, we have combined immunohistochemical staining for choline acetyltransferase with retrograde labeling of nucleus isthmi neurons after tectal injections of horseradish peroxidase. Seven fish received tectal horseradish peroxidase injections, and brain tissue from these animals was subsequently processed for the simultaneous visualization of horseradish peroxidase and choline acetyltransferase. In many nucleus isthmi neurons the dense horseradish peroxidase label obscured the choline acetyltransferase reaction product but horseradish peroxidase and choline acetyltransferase were colocalized in 54 cells from nine nuclei isthmi. The somata of nucleus reticularis mesencephali neurons stained so intensely for choline acetyltransferase that we could not determine whether they were labelled also with horseradish peroxidase. However, the large choline acetyltransferase-immunoreactive axons of nucleus reticularis mesencephali neurons stained intensely enough for us to follow them rostrally; the axons are clustered together until the level of the rostral tectum where two groupings form: one travels into the tectum and the other travels rostroventrally to cross the midline and enter the contralateral diencephalic preoptic area. We conclude therefore that cholinergic neurons project to the optic tectum from the nucleus isthmi as well as nucleus reticularis mesencephali in goldfish.     

The afferent connections of the tectum mesencephali in two chondrichthyans,the shark Scyliorhinus canicula and the ray Raja clavata

The afferent connection of the tectum mesencephali were studied in the spotted dogfish Scyliorhinus canicula and the thornback ray Raja clavata by means of the horseradish peroxidase (HRP) technique. Following unilateral injections in the tectum, labeled neurons could be identified in all main divisions of the brain and in the cervical spinal cord. Telencephalic neurons which project to the tectum mesencephali were observed in the caudal part of the pallium. Diencephalic projections to the tectum originate from the thalamus dorsalis pars medialis, the thalamus ventralis pars lateralis, the nucleus medius infundibuli, and the pretectal area. In Scyliorhinus labeled neurons could also be found in the corpus geniculatum laterale. Mesencephalic cells of origin of tectal afferent pathways were identified in the stratum cellulare externum of the contralateral tectum, in the nucleus tegmentalis lateralis, in the ventrolateral tegmentum, and in the nucleus ruber. Rhombencephalic cells projecting to the tectum could be identified in the nucleus cerebelli (only in Scyliorhinus), the nucleus vestibularis superior, the reticular formation, the nucleus funiculi lateralis, the nucleus tractus descendens nervi trigemini, and the nucleus dorsalis and intermedius areae octavolateralis. In addition a number of small-and medium-sized cells of the reticular formation were found labeled. Diffusely scattered labeled cells could be observed in the dorsal part of the cervical spinal cord. It is concluded that the tectal afferent connections in the chondrichthyans studied in general resemble those of other vertebrates, but that some striking differences exist. In particular, tectal afferents originating from the nucleus medius infundibuli, the nucleus cerebelli, and the nucleus dorsalis and intermedius areae octavolateralis have not been reported in other vertebrates.     

Afferents to the red nucleus in the lizard Podarcis hispanica: putative pathways for visuomotor integration.

The afferents to the red nucleus from visual and nonvisual forebrain centers have been investigated in the lizard Podarcis hispanica by using both retrograde and anterograde transport of tracers. Because the red nucleus constitutes a key structure in the limb premotor system, these sensory afferents probably are involved in visuomotor and other forms of sensorimotor integration. After tracer injections aimed at the red nucleus, retrograde labeling was found in the reticular thalamus, the subthalamus, the nucleus of the posterior commissure, as well as in two retinorecipient nuclei, namely, the ventral lateral and pretectal geniculate nuclei, where labeled cells are especially abundant. These geniculorubral projections have been confirmed by means of anterograde tracing with dextranamine injections. On the other hand, small injections of tracers in the retina demonstrated that its projections to the ventral lateral and pretectal geniculate nuclei are organized in a point-to-point fashion. Moreover, small tracer injections into the optic tectum of Podarcis indicated that the ventral lateral geniculate nucleus also receives a precisely organized tectal afferent. Taken together, these results strongly suggest that geniculorubral projections might constitute the neuroanatomical substrate for the generation of quick locomotor responses to appropriate visual stimuli. Additional ventral thalamic, subthalamic, and pretectal afferents to the red nucleus are likely to subserve other kinds of sensorimotor integration. These results help to clarify the organization of the reptilian motor system, including the telencephalic control of motor responses, and to unravel some of the major trends in the evolution of the limb premotor network of tetrapodian vertebrates.