Topography of the retinal projection to the superficial pretectal parvicellular nucleus of goldfish: a cobaltous-lysine study

The retinal projection to the superficial pretectal parvicellular nucleus (SPp) of goldfish was examined by filling select groups of optic axons with cobaltous-lysine. The tracer was applied intraocularly to peripheral retinal slits in some fish. In other fish, it was applied to optic axons from an intact hemiretina after one-half of the retina was ablated and the corresponding optic axons had degenerated. The results indicated that SPp is a folded structure, having a dorsal surface innervated by axons from temporal retinal ganglion cells and a ventral surface innervated by axons from nasal retinal ganglion cells. Peripheral retina innervates the anterodorsal and anteroventral edges of SPp, while central retina innervates the posterior genu. Dorsal retina innervates lateral SPp and ventral retina innervates medial SPp. Thus, although SPp is a folded nucleus, the topography of the retino-SPp projection is similar to the topography of the retinotectal projection. That is, the relative position of optic axons within SPp mirrors the retinal location of the ganglion cells that project to SPp. Retino-SPp axons occupy the center of the main optic tract before it divides into the two optic brachia. These axons are topographically arranged, with temporal retino-SPp axons being flanked on both sides by nasal retino-SPp axons. Retino-SPp axons arborize within SPp and then continue to enter the superficial tectal retino-recipient lamina. Thus, these axons innervate both SPp and the optic tectum. These findings are discussed with respect to chemospecific and morphogenetic views of visual system topography.     

Tectal projections in the goldfish (Carassius auratus): a degeneration study.

Efferents revealed by degeneration staining following tectal lesions in goldfish are presented. Four major projections were found. Ascending ipsilateral projections to pretectal-diencephalic areas exit the tectum rostrally and laterally and terminate in the area pretectalis (AP), lateral geniculate (LGN), nucleus pretectalis (NP), and nucleus rotundus (NR). Ascending contralateral projections exit rostrally and possibly laterally, enter the posterior and postoptic commissures and terminate in the contralateral AP, LGN, NP, NR, and rostral tectum. A medially directed projection enters the intertectal commissure, and some of these fibers may terminate sparsely in an area of the contralateral tectum homotopic to the lesion. A descending projection exits the tectum laterally and projects ipsilaterally to a dorsolateral tegmental nucleus (DLT) and the lateral reticular formation of the tegmentum and pons, and contralaterally to the medial reticular formation of the tegmentum and pons.     

Telencephalic projections to the goldfish hypothalamus: An anterograde degeneration study

In this study, large areas of goldfish telencephalon were ablated including rostral nucleus preopticus periventriculare (rNPP), and degenerating axons were traced by a modified Fink and Heimer procedure. The lesioning procedure ablated large regions of area dorsalis telencephali pars medialis, centralis, and dorsolateral complex; and completely removed area ventralis telencephali pars dorsalis, ventralis, and lateralis. In addition, the supracommissural nucleus and rNPP were lesioned specifically because both nuclei have been thought to be involved in courtship behavior and endocrine control of reproduction. This investigation demonstrated extensive fiber projections from telencephalic nuclei and/or rNPP to the hypothalamus. Lesioned telencephalon and/or rNPP projected bilaterally to nucleus preopticus and the suprachiasmatic nucleus and unilaterally to the following tuberal nuclei: nucleus anterior tuberis, and the lateral hypothalamic nucleus. A much larger fiber projection to the inferior lobe nuclei was also observed with a large contralateral as well as ipsilateral input.     

Retinal projections in the australian lungfish

Autoradiographic analysis of the primary retinofugal projections in the Australian lungfish reveals contralateral retinal projections to a ventral portion of the periventricular preoptic nucleus, throughout its rostrocaudal extent, and to 4 distinct terminal fields in the thalamus. Only one of these thalamic fields (t4) likely receives dendrites soley from dorsal thalamic neurons. Thalamic terminal field 1 probably receives dendrites from both dorsal and ventral thalamic neurons, and fields 2 and 3 from only ventral thalamic neurons. Contralateral retinofugal fibers terminate in the pretectum and in the superficial and central tectal zones. The central tectal terminal field is restricted to the medial one-third of the tectum. At pretectal levels a contralateral basal optic tract arises from the marginal optic tract and terminates along the lateral edge of the tegmentum, as a series of glomerular puffs, and in the rostral pole of a superficial isthmal nucleus. The Australian lungfish, unlike the African and South American lungfish, possesses ipsilateral retinal projections to all of the nuclei that receive contralateral retinal input.     

Retinal projections in gymnotid fishes

Retinal projections were studied in four species of gymnotid fishes, Gymnotus carapo, Hypopomus artedi, Eigenmannia virescens and Sternopygus sp. with the aid of cobalt or HRP labelling and autoradiographic techniques. The optic tract gives off a small branch, the axial optic tract and then, after crossing in the midline, splits into a dorsomedial, dorsal and ventral fascicle. E. virescens and Sternopygus sp. display in addition an accessory optic tract. In all four species retinal projections are bilateral; ipsilateral projections, however, are extremely sparse. In all four species, the retinal fibres terminate bilaterally in the suprachiasmatic nucleus, dorsolateral optic nucleus of the thalamus and the optic nucleus of the posterior commissure; a bilateral retinotectal projection was only found in E. virescens and G. carapo. Retinal projections are only contralateral to the ventromedial nucleus of the thalamus, the central pretectal nucleus and the accessory optic nucleus. The contralateral retinotectal fibres terminate in the stratum fibrosum and griseum superficiale, and in the stratum album centrale and stratum periventriculare. A small accessory optic tract and nucleus were detected in E. virescens and Sternopygus sp. but not in G. carapo and H. artedi. The results indicate that the visual system of gymnotid fish is as simple as that of mormyrids. The poor visibility in the environment where these animals live and the additional sensory system which these animals possess may explain the poor development of the visual system.     

Retinal projections in the lizard Podarcis hispanica

The retinofugal of the lizard Podarcis hispanica has been examined by means of enzymatic method with horseradish peroxidase (HRP). The retinal ganglion cells project contralaterally to thalamus (nucleus geniculatus lateralis pars dorsalis, nucleus geniculatus lateralis pars ventralis and nucleus ventrolateralis pars ventralis and nucleus ventrolateralis pars ventralis), pretectum (nucleus lentiformis mesencephali, nucleus geniculatus pretectalis and nucleus posterodorsalis) and optic tectum (layers 14 and 12, mainly, and layers 13 and 11). A small ipsilateral tract has been observed. Some of these fibers project to the lateral geniculate complex and the nucleus ventrolateralis pars ventralis. Most of the ipsilateral fibers have been observed in the neuropil of nucleus geniculatus pretectalis and the layer 14 of the optic tectum. The ipsilateral component, an inconstant structure in reptiles, presents an important development in Podarcis hispanica, although the number of its fibers is relatively small.     

Retinal terminals in the goldfish optic tectum: Identification and characterization

Retinal terminal profiles in the goldfish optic tectum were identified electron microscopically after (1) labeling with horseradish peroxidase and (2) in the early stages of degeneration in short-term eye enucleates. All labeled terminals shared certain common morphological characteristics which were identical to those of a population of terminals in normal tecta. Terminals of this type disappeared 30 days after enucleation of the contralateral eye. Retinal terminal presynaptic profiles were characterized by (1) round and oval synaptic vesicles; (2) mitochondria with irregular, randomly oriented cristae, large intracristal spaces, dilated membrane spaces, and primarily light matrices; (3) a wide range in profile area, 0.06–6.82 μm²; (4) large numbers of synaptic vesicles per profile area 168± 33 synaptic vesicles per μm²; (5) asymmetric synapses; and (6) multiple synaptic contacts (1.46 ± 0.73 per terminal profile). The postsynaptic elements included both dendritic and, less commonly, pleomorphic vesicle-containing profiles. The majority of postsynaptic dendritic profiles were small (0.01–0.40 μm²). Serial synaptic contacts were occasionally seen. The combination of vesicular and mitochondrial morphology (1 and 2 above) was necessary and sufficient to establish the retinal origin of a terminal, but use of such criteria would underestimate the number of retinotectal terminals by omitting those which did not have a mitochondrion in the plane of section. The number of such terminals was calculated from independent measurements, and the total number of retinal terminal profiles per area of neuropil was estimated.     

Retinal synaptic arrays: continuing development in the adult goldfish.

We report a light- and electron-microscopic examination of the inner plexiform layer of the central retina of young (c. 1 year) and old (3-4 year) goldfish. There were no new neurons added to this region during the growth period. Nonetheless, there were substantially more synapses (per cell, per mm2, or per degree 2) in the older retinas. This result is discussed in the contexts of retina function and neural development.     

Retinal projections in tyrosinase-negative albino cats

Retinal projections were examined in two tyrosinase-negative albino cats using autoradiographic techniques. Cats from this colony have pink eyes; their retinal pigment epithelium, ciliary body, and iris epithelium are completely devoid of melanin pigment. Test breeding for five generations indicates that these cats are true albinos (cc). The most striking feature of the albino cats' visual pathways was a reduction in ipsilateral input which was more severe than that reported for Siamese cats. The only evidence of ipsilateral input to the laminated dorsal lateral geniculate nucleus of the albino was a small lateral normal segment and a small projection to the lateral portion of lamina C1. Ipsilateral projections to the medial interlaminar nucleus, retinal recipient zone of the pulvinar complex, ventral lateral geniculate nucleus, superior colliculus, and pretectum also were reduced. Ipsilateral projections to the suprachiasmatic nucleus appeared to be normal in the albino cats studied. Our findings indicate that, compared to the normal cat as well as to the tyrosine-positive Siamese cat, the tyrosinase-negative albino has reduced ipsilateral retinal projections. The albino cat is a model system analogous to tyrosinase-negative albinism found in mammals.     

Retinal projections in the ringtailed possum Pseudocheirus peregrinus.

The retinal projections in the ringtailed possum, Pseudocheirus peregrinus were determined using Fink-Heimer material and autoradiography. At least seven regions in the brain receive retinal projections. These are (1) the suprachiasmatic nucleus of the hypothalamus (2) the dorsal lateral geniculate nucleus (3) the ventral lateral geniculate nucleus (4) the lateral posterior nucleus (5) the pretectum (6) the superior colliculus, and (7) the accessory optic system. The accessory optic system and lateral posterior nucleus receive a contralateral retinal projection only and the other five regions receive a bilateral retinal projection. The dorsal lateral geniculate nucleus consists of two parts: an outer alpha division of closely packed cells and an inner beta division containing loosely scattered cells. There are no cell layers apparent within the alpha division in Nissl sections. The autoradiographs and Fink-Heimer material reveal four concealed laminae within the alpha division. Lamina 1, which is adjacent to the optic tract and lamina 3 receive a predominantly contralateral input. Laminae 2 and 4 receive a predominantly ipsilateral input. The beta segment contains a fifth lamina which receives contralateral retinal input.     

Retinal projections to the pregeniculate nucleus in the hemispherectomized monkey

Intraocular injections of tritiated proline were used to test the hypothesis that unilateral removal of all visual cortical areas results in increased distribution of retinal terminals in the pregeniculate nucleus (PGN) of the thalamus in monkeys. Following hemispherectomy, retinal input to the ipsilateral PGN was reduced by an average of 18.5% when compared to its contralateral homologue, which corresponded to the reduction in nuclear volume (19.3%). Our results show that removal of cortical afferents to the external layer of the PGN does not induce invasion of retinal projections into this region of the nucleus.     

Retinal origin of the fasciculus medialis tractus optici in adult goldfish

In teleosts, the fasciculus medialis tractus optici departs the optic tract and follows an aberrant path towards the midbrain tectum. To determine the retinal origin of the fascicle, horseradish peroxidase was injected into localized lesions of either the optic nerve or the retina. The fasciculus medialis axons of adult goldfish derive selectively from ganglion cells residing in the dorsotemporal portion of the central retina. The fasciculus medialis appears to be an integral part of the chronotopic ordering of fibers in the optic tract.     

Retinal projections in adult and newborn grey squirrels

Retinal projections in newborn squirrels were compared to those in adults by using horseradish peroxidase (HRP) as a highly sensitive anterograde tracer. In both newborn and adult squirrels, the HRP reaction product was found in the dorsal lateral geniculate nucleus, the superior colliculus, the pretectal nuclei, and the nuclei of the accessory optic tract. Thus, newborn squirrels have retinal input to most or all structures normally innervated in the adult. However, the pattern of terminations differed in the newborn from that in the adult, and this was especially apparent in the dorsal lateral geniculate nucleus and the superior colliculus. In the dorsal lateral geniculate nucleus, the regions of ipsilateral and contralateral retinal inputs were clearly less segregated than in adults, although the adult laminar pattern of retinal terminations was partially apparent, even though there was yet no cytoarchitectural evidence of the adult lamination pattern. In the superior colliculus, a marked difference was seen in the pattern of ipsilateral retinal terminations. In the adult, ipsilateral retinotectal input was restricted to a narrow, dense, patchy, mediolateral band in stratum opticum in the rostral colliculus. In the newborn, the ipsilateral retinotectal input was less dense, free of patches, spread in thickness to include much of the stratum opticum and the superficial grey, and spread in extent to include all but the caudal pole of the colliculus. These observations are consistent with the prevailing view that visual connections are initially widespread and become restricted during the course of development.     

Retinal projections in lamprey (Lampetra fluviatilis)

Visual projections in lamprey were investigated using two methods,--one by revealing transport of horseradish peroxidase, and the other by silver impregnation of degenerating axons and terminals after enucleation of the eye. Both methods produced similar results. The chiasm showed incomplete crossing of retinal fibres, the major part of which, as an optic tract, proceed along the contralateral thalamus up to the entry into the optic tectum, while the smaller part takes the same course on the ipsilateral side. Besides, from the posterior part of the optic chiasm an axial optic tract branches off, which proceeds through the central part of the contralateral thalamus up to the pretectal nucleus, individual fibres of which enter the central grey layer of the optic tectum. On the contralateral side, the visual projections are localized in the lateral geniculate body, pretectal nucleus, in the three upper layers of the optic tectum, in the ventrolateral area of the optic tectum and as solitary diffuse projections in the mesencephalic tegmentum. Innervation of thalamic and pretectal nuclei are realized by two tracts--the tractus opticus proper, and tractus opticus axialis. On the ipsilateral side visual projections, excepting the optic tract, are scarce and in the thalamus appear as small areas of the lateral geniculate body and pretectum adjacent to the optic tract. Solitary visual projections were found in two upper layers of the rostral optic tectum and in larger numbers in the 3rd and 4th layers of the caudal part and in ventrolateral area of the optic tectum. Projections in mesencephalic tegmentum were single. Diffuse visual projections in the lateral part of hypothalamus could be revealed only by the silver impregnation method. Using the peroxidase method two types of cells were observed in mesencephalic tegmentum where, possibly, the centrifugal fibres proceeding to retina, originate. A comparison is made of central visual projections in lampreys and other representatives of nonmammalian vertebrates.     

Visual orienting response in goldfish: a multidisciplinary study

The neural basis underlying the orienting response has been thoroughly studied in frontal-eyed mammals. However, in non-mammalian species, including fish, it remains almost unknown. Therefore, we studied the contribution of the optic tectum and the mesencephalic reticular formation to the performance of the orienting response in goldfish, using behavioural, physiological, and anatomical tracer techniques. The appearance of a visual stimulus (a pellet of food) in the environment of a goldfish evoked a turn of the body to reorient the line of sight. Left-tectal lobe ablation abolished the orienting turn response towards the contralateral hemifield. Electrical microstimulation of the optic tectum suggested the presence of a motor map, which is in correspondence with the overlying visual representation, as previously reported in other vertebrates. The tracer biotin-dextran amine was injected into different functionally identified tectal zones. The results showed that rostral and caudal poles of the mesencephalic reticular formation receive outflow mainly from the rostral and caudal tectal poles, respectively. This suggests that the tectal wiring with downstream structures is site-dependent. Furthermore, the electrical activation of rostral and caudal mesencephalic reticular formation revealed a different contribution to vertical and horizontal orienting eye movements. We conclude that the basic neural system coding the orienting response appears early in phylogenesis, although some specific characteristics are selected by adaptive pressure.     

Retinal projections in the Tasmanian devil, Sarcophilus harrisii.

Retinal projections were mapped in Tasmanian devils which had one eye injected with 3H-proline. The retinal fibers terminate in seven regions in the brain. These are (1) dorsal lateral geniculate nucleus (LGNd), (2) ventral lateral geniculate nucleus, (3) lateral posterior nucleus, (4) pretectum, (5) superior colliculus, (6) hypothalamus and (7) accessory optic system. The pattern of retinal input to six of these regions is similar to that seen in other marsupials. The pattern of retinal projections to the LGNd, while basically similar to that observed in other polyprotodont marsupials, is much simpler than that seen in the related native cat, Dasyurus viverrinus. The LGNd of Sarcophilus presents the simplest cytoarchitectural organisation of any marsupial examined so far. Each LGNd receives overlapping projections from both eyes. Suggestions of an intermittent lamination are seen in the LGNd contralateral to an eye injection of 3H-proline. On the ipsilateral side there are two patches of label, a large lateral patch and a smaller medial patch, both of which occupy areas receiving contralateral input. The monocular segment, occupying the ventral 40% of the nucleus, is more extensive than has been reported in any other polyprotodont marsupial.