Retinotopic analysis of fiber pathways is amphibians. I. The adult newt Cynops pyrrhogaster

The possibility that pathways of retinal fibers within the optic tract and the tectum of the adult newt are retinotopic was examined by selective labeling of the retinal fibers with horseradish peroxidase. Within the optic tract fibers from the ventral, temporal and dorsal retinal quadrants were ordered from the dorsal to ventral edges of th optic tract. The nasal retinal fibers exhibited two different pathways. The fibers from the dorsonasal retina ran along the ventral edge of the optic tract, while the fibers from the ventronasal retina ran along the dorsal edge of the optic tract. Segregation of pathways within the optic tract was incomplete between the nasal and other retinal fibers. The dorsonasal retinal fibers were mixed completely with the dorsal retinal fibers, and the ventronasal retinal fibers were mixed partly with the ventral retinal fibers. Both the dorsal and dorsonasal retinal fibers preferentially entered the lateral tract, and finally projected onto the ventrolateral parts of the middle tectum and of the caudal tectum, respectively. The ventral and ventronasal retinal fibers entered the dorsomedial tract, and projected onto the dorsomedial parts of the middle tectum and of the caudal tectum, respectively. The temporal retinal fibers invaded the nasal tectum directly. Most dorsal, ventral, and nasal retinal fibers ran along the sub-tracts as far as to the level of their terminals, then sharply turned in a direction to the tectum.     

Retinotopic analysis of fiber pathways in amphibians. I. The adult newt Cynops pyrrhogaster

The possibility that pathways of retinal fibers within the optic tract and the tectum of the adult newt are retinotopic was examined by selective labeling of the retinal fibers with horseradish peroxidase.Within the optic tract fibers from the ventral, temporal and dorsal retinal quadrants were ordered from the dorsal to ventral edges of the optic tract. The nasal retinal fibers exhibited two different pathways. The fibers from the dorsonasal retina ran along the ventral edge of the optic tract, while the fibers from the ventronasal retina ran along the dorsal edge of the optic tract. Segregation of pathways within the optic tract was incomplete between the nasal and other retinal fibers. The dorsonasal retinal fibers were mixed completely with the dorsal retinal fibers, and the ventronasal retinal fibers were mixed partly with the ventral retinal fibers.Both the dorsal and dorsonasal retinal fibers preferentially entered the lateral tract, and finally projected onto the ventrolateral parts of the middle tectum and of the caudal tectum, respectively. The ventral and ventronasal retinal fibers entered the dorsomedial tract, and projected onto the dorsomedial parts of the middle tectum and of the caudal tectum, respectively. The temporal retinal fibers invaded the nasal tectum directly.Most dorsal, ventral, and nasal retinal fibers ran along the sub-tracts as far as to the level of their terminals, then sharply turned in a direction toward the tectum.     

Retinotopic analysis of fiber pathways in amphibians. II. The frog Rana nigromaculata

The possibility of retinotopic organization of pathways of retinal fibers within the optic tract and the tectum of the frog was studied by selective labeling of the retinal fibers with horseradish peroxidase. Within the optic tract the pathways of the ventral, temporal and dorsal retinal fibers were ordered from the dorsal to ventral edges of the optic tract. The nasal retinal fibers ran along both the dorsal and ventral edges of the optic tract. The dorsal retinal fibers and the nasal retinal fibers which were located along the ventral edge of the optic tract entered the ventrolateral perimeter of the tectum and formed the lateral tract. The ventral retinal fibers and the nasal retinal fibers which were located along the dorsal edge of the optic tract entered the dorsomedial perimeter of the tectum and formed the dorsomedial tract. The temporal retinal fibers invaded the tectum directly at the diencephalo-tectal junction. The topography of fiber pathway observed for the frog was exactly the same as that seen in the newt, and seemed to be common to all amphibian species.     

A retinotopic analysis of the central connections of the optic nerve in the frog

The possibility of retinotopic organization in the optic nerve projections to the contralateral and ipsilateral diencephalon was studied by means of partial retinal lesions and staining for terminal degeneration by the Fink-Heimer technique. A retinotopic pattern of projection was observed in the nucleus of Bellonci, the corpus geniculatum thalamicum and the posterior thalamic nucleus. The temporal quadrant of the retina, and, to a lesser extent, the ventral quadrant projected to the ipsilateral side as well as to the contralateral side. In each diencephalic region noted above, the temporal and dorsal quadrants of the retina were represented more posteriorly (posteroventrally), and ventral and nasal quadrants projected more anteriorly (anterodorsally). The areas of representation for the temporal and ventral quadrants were located superior (superoposterior) to those for the dorsal and nasal quadrants. In their overall configuration and orientation, the retino-diencephalic maps show mirror-image reversal with respect to the retino-tectal projection. Since, in their areal extent, both the retino-diencephalic maps and the retino-tectal map are approximately parallel to the ventricular surface, their mirror-image reversal appears to indicate a reversal in the polarity of developmental processes across the di-mesencephalic junction. The retinotopic organization within the optic tract in the diencephalon and tectum was also analyzed. In the optic tract, the quadrants of the retina are reassembled such that the dorsal and nasal quadrants are widely separated in, respectively, the ventral and dorsal edges of the tract; the temporal and ventral quadrants are systematically represented in intermediate levels in the tract, the temporal quadrant above the dorsal, and the ventral quadrant below the nasal. When the optic tract bifurcates to encircle the tectum, the fibers from the ventral and nasal quadrants enter the dorsomedial arm and the fibers from the temporal and dorsal quadrants enter the ventrolateral arm of the optic tract. The paths taken by optic fibers in traversing the tectum to reach their areas of termination were reconstructed. Many optic fibers show an alignment parallel to an anteroventral posterodorsal axis as they cross the surface of the tectum, but the OS vs IS characterization of the fibroarchitecture of the tectum appears to be an oversimplification.     

Projections of growth-cone-bearing fibers of retinal ganglion cells within co-cultured tectal explants: early branching depends on age of target tissue

The projections of retinal ganglion cell axons within co-cultured tectal explants were analyzed in order to investigate some of the factors that determine the earliest responses of retinal axons to cues present in an isolated target tissue. Half retinas and superior colliculi (tecta) from the embryonal mouse were explanted, separated by a 0.5 mm gap. After 5 days in vitro retinal ganglion cells were labeled by extracellular ionophoresis of HRP into the optic nerve head region. Cleared co-cultures were studied as whole mounts. Growth-cone-bearing retinal fibers were studied in standard tectal co-cultures, and in cases where tectum had been explanted 2 weeks prior to retina. The heterochronously prepared cultures had a higher proportion of fibers with complex branching patterns than the synchronous explants. Cultures in which retinas were explanted 1 week after tecta exhibited intermediate proportions of such fibers. These observations suggest that older tecta facilitate branching of ingrowing retinal fibers, although other alterations during in vitro development must be evaluated. The growth patterns of axons originating in nasal and temporal hemi-retinas were analyzed in terms of possible positional cues provided by the target tecta. Axons originating in temporal hemi-retinas did not show evidence of preferential branching in, or growth toward, appropriate anterior regions of co-cultured tectal explants. In contrast, the majority of nasal retinal axons showed enhanced terminations and complex branching in, and bending towards, the posterior tectum.     

The optic system of the teleost Cichlasoma biocellatum

The retinal projections of the fresh water teleost. Cichlasoma biocellatum, were examined using a modification of the Nauta-Gygax method following unilateral enucleations and removal of retinal quadrants. After the unilateral enucleations, degeneration was found in the fasciculus medialis nervioptici, fasciculus geniculatus tractus optici, fasciculus dorsomedialis tractus optici, and the accessory optic tract. The pretectal nucleus, nucleus corticalis, and accessory optic nucleus contained debris of terminal degeneration. The majority of the optic fibers terminated in five layers in the optic tectum. The quadrant removals showed that fibers from the retinal quadrants formed layers in the ribbon shaped optic nerve running the width of the nerve. The dorsal to ventral arrangement of the fibers was: dorsal nasal quadrant; dorsal temporal quadrant; ventral nasal quadrant; and the ventral temporal quadrant. The fibers from only the posterior half of the retina supplied the fasciculus geniculatus tractus optici and the corpus geniculatum laterale. Fibers from the anterior half of the retina supplied the accessory optic tract and nucleus. All other tracts and nuclei examined received fibers from all the retinal quadrants. The projection of the retinal quadrants to the tectal quadrants was specific and always formed five layers of degeneration.     

Retinotopic organization of the developing retinotectal projection in the zebrafish embryo

Developing retinal axons in the zebrafish embryo were stained with HRP or with the fluorescent dyes dil and diO to study the formation of the retinotectal projection. Retinal axons leave the eye at 34-36 hr postfertilization (PF), invade the tectum at 46-48 hr PF, and innervate the tectal neuropil at 70-72 hr PF. Dorsal and ventral axons occupy separate aspects of the optic nerve and tract and pass into their retinotopically appropriate ventral and dorsal hemitectum, respectively. Nasal and temporal axons are segregated in the nerve, mixed in the tract, and are coextensive over the rostral half of tectum until 56 hr PF. They then segregate again, due to the progression of nasal axons into the open caudal tectum. Thus, at 70-72 hr PF, dorsal and ventral as well as temporal and nasal axons occupy their retinotopically appropriate tectal quadrants. After ablation of the temporal retina prior to the time of axonal outgrowth, the nasal axons bypass the vacant rostral tectum to terminate in the caudal tectal half. Temporal axons in the absence of nasal axons remain restricted to their appropriate rostral tectal half, suggesting that nasal and temporal axons possess a preference for their retinotopically appropriate tectal domains. Measurements of individual terminal arbors and the tectal areas in embryos and in adult zebrafish showed that individual arbors are large with respect to the embryonic tectum but are about 14-15 times smaller than in the adult. However, the proportion of tectum covered by embryonic arbors is about 7 times larger than in the adult, suggesting that a higher precision of the adult projection is achieved as a result of a greater enlargement of the tectum than of the arbors.     

The course of axons of retinal ganglion cells within the optic nerve and tract of the chick (Gallus gallus)

Small laser lesions placed in the posthatch chicken retina resulted in axotomy and then death of all ganglion cells located in a sector peripheral to the primary damage. With the use of silver techniques, the patterns of degenerating retinal fibers in the optic nerve, chiasm, and optic tract were examined. In the proximal part of the optic nerve, radial retinal lesions resulted in a sheet of degenerating axons along the rostrocaudal extent of the nerve. The position of degenerating axons was related to the site of their entry in the optic nerve head with an overlapping distribution of degenerating fibers entering the optic nerve head from equivalent points from the temporal and nasal sides. In the optic chiasm, the distribution of fibers was similar to that seen in the proximal part of the optic nerve. In the optic tract there was a similar mixing of fibers from opposite sides of the retina. The ventral, nasal and temporal retinal fibers lay in the superficial part of the tract whereas the fibers from the nasal and temporal dorsal retina ran in the deeper, medial aspect of the tract. The central-to-peripheral axes of the retina were mapped along the rostrocaudal axis of the tract. As the tract approached the tectum degenerating fibers from single retinal lesions did not always remain together. In the case of a lesion in the ventral nasal retina, degenerating fibers split into two bundles located at opposite ends of the tract only to reunite at their terminal regional at the caudal pole of the tectum.(ABSTRACT TRUNCATED AT 250 WORDS)     

Retinal projections and retinal ganglion cell distribution patterns in a sturgeon (Acipenser transmontanus), a non-teleost actinopterygian fish

Retinal projections in a sturgeon were studied by injecting biocytin or HRP into the optic nerve. The target areas are the preoptic area, thalamus, area pretectalis, nucleus of posterior commissure, optic tectum, and nuclei of the accessory optic tract. Furthermore, a few labeled fibers and terminals were found in a ventrolateral area of the caudal telencephalon. All retinal projections are bilateral, although contralateral projections were more heavily labeled. Retrogradely labeled neurons were found in the ventral thalamus bilaterally. Retinal projections in sturgeons are similar to those of other non-teleost actinopterygians and chondrichthyans (sharks), in terms of the targets and extent of bilateral projections. Distribution patterns of ganglion cells in the retina were examined in Nissl-stained retinal whole mount preparations. The highest density areas were found in the temporal and nasal retinas, and a dense band of ganglion cells was observed along the horizontal axis between the nasal and temporal areas of highest density. The density of ganglion cells in the dorsal retina is the lowest. The total number of ganglion cells was estimated to be about 5 x 10(4) in a retina. The existence of a low density area in the dorsal retina suggests reduced visual acuity in the ventral visual field.     

Retinotopic analysis of fiber pathways in the regenerating retinotectal system of the adult newt cynops pyrrhogaster

Retinotopic analysis of the pathways of regenerating retinal fibers within the optic tract and in the tectum of an adult newt was performed by selective labeling of the retinal fibers with horseradish peroxidase. At the tenth week of regeneration, all the regenerating retinal fibers from different retinal quadrants had terminal arbors nearly at the parts of the tectum innervated normally by those quadrants. The pathways for individual retinal fibers, however, were greatly disorganized within the optic tract and did not show any retinotopic ordered geography. The most rostral segregation of pathways of regenerating fibers was observed at the diencephalo-tectal junction. THe temporal retinal fibers invaded the tectum directly, while the dorsal, ventral and nasal retinal fibers generally shifted toward the dorsomedial or the lateral direction, as if they traced the dorsomedial or the lateral tracts formed in normal newt. The direction of the shifting of fiber pathways, however, did not depend on the origins of retinal fibers within retinal circumference, but depended on the location of fibers with in the optic tract. As a result, a large number of regenerating fibers reached their normal sites of innervation within the tectum via anomalous routes. These mis-routed fibers did not form branches or terminal arbors at ectopic parts within the tectum.     

Spatial organization of the retinal projection to the avian lentiform nucleus of the mesencephalon

Utilizing the horseradish peroxidase retrograde tracing technique and the 2-deoxy-D-glucose metabolic mapping technique, we have demonstrated in chickens the distribution of retinal ganglion cells that project to the lentiform nucleus of the mesencephalon (LM) and the retinotopic organization of the projection in the LM. Retinal ganglion cells labeled after a nearly complete injection into the LM were found in the four quadrants, distributed in a wide horizontal belt lying along both sides of the retinal equator and stretching from the temporal to the nasal retina. The HRP-labeled cells, which appeared round or oval, ranged from 25 to 840 micron 2 in size with most in the smaller size range. Results of partial HRP injections into the LM and metabolic mapping patterns in the LM produced by stimulation of half the retina with horizontal visual motion suggest that there is an orderly mapping of the retina onto the LM. The inferior temporal quadrant projects to the rostrodorsal LM; the inferior nasal quadrant projects to the caudodorsal LM. The superior temporal quadrant projects to the middle and ventral LM, extending from the rostral to the caudal pole, whereas the superior nasal quadrant projects to a small zone in the caudal LM. The mapping of the retinal quadrants in the LM is remarkably similar to that reported in the optic tectum of birds. We suggest that a common embryological anlage with the optic tectum and the arrangement of retinal axons in the optic tract are important factors in establishing the retinotopic organization of the LM.     

Retinal fibers alter tectal positional markers during the expansion of the retinal projection in goldfish

Although widely accepted, the theory, that neurones carry immutable cytochemical markers which specify their synaptic connections, is not consistent with plastic reorganizations. Half retinal fish were therefore tested for changed markers following expansion. Optic nerve crush at the time of the half retinal ablation resulted in regeneration of a normal, restricted projection; but nerve crush following expansion (many months later) resulted in reestablishment of the expanded projection, assessed both by electrophysiological mapping and by radioautography. Since this implied changed markers, the half retina and tectum were tested independently using the ipsilateral tectum and eye as controls. In normal fish, removal of one tectum and deflection of the corresponding optic tract toward the remaining tectum resulted in regeneration of a positionally normal but ipsilateral map. In experimental fish, after the half retina had expanded its projection to the contralateral tectum, its optic tract was deflected to the control tectum. After 40 days it had regenerated a normal, restricted map indicating that the retinal markers had not changed. Such restricted projections did not expand in the presence of the normal projection even after a year or more. Similarly, the optic tract from the normal eye was deflected to cause innervation of the tectum containing the expanded half retinal projection. After 40 days, the projection regenerated from the normal eye was similar to the expanded half retinal projection. Areas of the normal retina corresponding to the missing areas of the half retina were not represented. Tectal markers had been altered by the half retinal fibers. In a final group, tecta were denervated and tested at various intervals by innervation from ipsilateral half retinal eyes. After five months of denervation, the regenerating fibers were no longer restricted to the rostral tectum but formed an expanded projection initially. Apparently tectal markers are induced by the retinal fibers, changed during expansion, and disappear during long-term denervation.     

Gradients of ephrin-A2 and ephrin-A5b mRNA during retinotopic regeneration of the optic projection in adult zebrafish

Regeneration of optic axons in the continuously growing optic system of adult zebrafish was analyzed by anterograde tracing and correlated with the mRNA expression patterns of the recognition molecules ephrin-A2 and ephrin-A5b in retinal targets. The optic tectum and diencephalic targets are all reinnervated after a lesion. However, the rate of erroneous pathway choices was increased at the chiasm and the bifurcation between the ventral and dorsal brachium of the optic tract compared to unlesioned animals. Tracer application to different retinal positions revealed retinotopic reinnervation of the tectum within 4 weeks after the lesion. In situ hybridization analysis indicated the presence of rostral-low to caudal-high gradients of ephrin-A2 and ephrin-A5b mRNAs in unlesioned control tecta and after a unilateral optic nerve lesion. By contrast, the parvocellular superficial pretectal nucleus showed retinotopic organization of optic fibers but no detectable expression of ephrin-A2 and ephrin-A5b mRNAs. However, a row of cells delineating the terminal field of optic fibers in the dorsal part of the periventricular pretectal nucleus was intensely labeled for ephrin-A5b mRNA and may thus provide a stop signal for ingrowing axons. Ephrin-A2 and ephrin-A5b mRNAs were not detectable in the adult retina, despite their prominent expression during development. Thus, given a complementary receptor system in retinal ganglion cells, expression of ephrin-A2 and ephrin-A5b in primary targets of optic fibers in adult zebrafish may contribute to guidance of optic axons that are continuously added to the adult projection and of regenerating axons after optic nerve lesion.     

Tectotectal connectivity in goldfish.

The vertebrate optic tectum is a functionally coupled bilateral structure which plays a major role in the generation of motor commands for orienting responses. However, the characteristics of the tectotectal connectivity are unknown in fish, and have been reported only to a limited extent in other vertebrates. The purpose of the present study was to determine the anatomical basis underlying the functional coupling between tecta in goldfish, and to identify both similarities and differences to those features reported in other vertebrate species. The present experiments used the bidirectional tracer biotinylated dextran amine to map the distribution of labeled cells and synaptic boutons in the contralateral tectum following injections into identified tectal sites. Fibers that interconnect both tecta coursed through the tectal commissure. The cells of origin of these fibers, the tectotectal cells, and their synaptic endings were located in the deep layers, mainly in the strata periventricular and griseum central, respectively. Corresponding sites throughout the two tecta were interconnected in a symmetrical point-to-point fashion. The tectal commissure was composed of at least two distinct bundles of axons, which differed in their dorsoventral location, fiber diameter, and projection targets. The dorsal axons were tectotectal axons, they were thinner in diameter and profusely branched, and gave off en passant and terminal boutons in the deep layers of the contralateral tectum. The ventral axons were thicker in diameter, and formed the contralateral tectofugal-descending tract. Such fibers had few axon collaterals and boutons in the contralateral tectum. Boutons adjacent to retrogradely labeled tectotectal cells were very scarce. The data are discussed in terms of the coupling between tecta generating the motor commands required for orienting movements.     

Relationship of ocular pigmentation to the boundaries of dorsal and ventral retina in a nonmammalian vertebrate

The goldfish eye and retina are partitioned traditionally into dorsal and ventral sectors by a horizontal meridian that passes through the optic disc and is perpendicular to a vertical meridian that extends from the remnant of the choroid fissure through the optic disc. Axons of retinal ganglion cells (RGCs) situated above the horizontal meridian are thought to reach the optic tectum via the ventrolateral optic tract and axons of RGCs situated below the horizontal meridian are thought to reach the optic tectum via the dorsomedial optic tract. When cobaltous-lysine was applied to small temporal retinal slits that were centered on the traditional horizontal meridian, filled fibers were found in the dorsomedial, but not in the ventrolateral, optic tract (Springer and Mednick, '83). Since cobalt-filled axons should have been found in both optic tracts, the traditional horizontal meridian does not indicate the actual boundary between dorsal and ventral retina. We report here that the goldfish iris contains nasal and temporal pigmentation lines (darts) that are each located approximately 21 degrees above the traditional horizontal retinal meridian. Cobalt applied to retinal slits located just above the darts filled RGC axons in the ventrolateral optic tract and cobalt applied to retinal slits just below the darts filled RGC axons in the dorsomedial optic tract. Converging evidence for the reliability of the darts as indicators of the boundary between dorsal and ventral retina was obtained by applying cobalt to severed RGC axons along the dorsomedial edge of the tectum. Cobalt-filled RGCs were found below the nasal dart.(ABSTRACT TRUNCATED AT 250 WORDS)     

The organization of the fibers in the optic nerve of normal and tectum-less Rana pipiens

We have examined the detailed order of retinal ganglion cell (RGC) axons in the optic nerve and tract of the frog, Ranapipiens. By using horseradish peroxidase (HRP) injections into small regions of theretina, the tectum, and at various points along the visual pathway, it hasbeen possible to follow labelled fibers throughout their course in the nerve and tract. Several surprising features in the order of fibers in the visual pathway were discovered in our investigation. The fascicular pattern of RGC axons in che retina is similar to that described in other vertebrates; however, immediately central to their entry into the optic nerve head, approximately half of the fibers from the nasal or temporal retina cross over to the opposite side of the nerve. Although the axons from the dorsal and ventral regions of the retina generally remain in the dorsal and ventral regions of the nerve, some fiber crossing occurs in those axons as well. The result of this seemingly complex rearrangement is that the optic nerve of Rana pipiens contains mirror symmetric representations of the retinal surface on either side of the dorsal ventral midline of the nerve. The fibers in each of these representation are arranged as semicircles representing the full circumference of the retina. This precise fiber order is preserved in the nerve until immediately periphearal to the optic chiasm, at which point age-related axon from both side of the nerve bundle together. Consequently, when a small pellet of HRP is placed in the chiasmic region of the nerve, an annualus of retinal ganglion cells and a corresponding annulus of RGC terminals in the tectum are la belled. As the age-related bundles of fibers emerge from the chiasm they split to form a medial bundle and a lateral bundle, which grow in the medial and lateral branches of the optic tract, respectively. Although the course followed by RGC axons in the visual pathv/ay is complex, we propose a model in which the organization of fibers in the nerve and tract can arise from a few rules of axon guidance. To determine whether the optic tecta, the primary retinal targets, play a role in the development and organization of the optic nerve and tract, we removed the tectal primordia in Rana embryos and examined the order in the nerve when the animals had reached larval stages. We found that the order in the nerve and tract was well preserved in tectumless frogs. Therefore, we propose that guidance factors independent of the target direct axon growth in the frog visual system.     

Mapping the normal and regenerating retinotectal projection of goldfish with autoradiographic methods

In normal goldfish, lesions of various size were made in nasal or temporal retina immediately prior to retinal labeling with tritiated proline. The resulting gaps in retinal innervation of tectum indicated that the projection is retinotopographically ordered to a precision of about 50 μm. Similarly, acute tectal incisions transecting the optic pathways were combined with immediate retinal labeling. The resulting tectal denervation confirmed that most fibers follow highly ordered paths through the stratum opticum of tectum; but a few fibers were found to follow unusual paths to their appropriate tectal positions. In other fish, the optic nerve was crushed. At various times afterwards, retinotopography and pathway order were similarly analyzed by making retinal lesions or tectal incisions just prior to labeling. For up to 40 days after crush, the projection lacked any refined retinotopic order. Only a gross topography could be demonstrated. Over several months, retinotopography gradually improved eventually approaching that of normals. Correlated with this was an initial stereotypic growth through the pathways of the stratum opticum followed by a long period of highly anomalous growth through the innervation layer. Evidently, many regenerated fibers grew in through inappropriate routes to the wrong region of tectum but subsequently arrived at their appropriate locus by circuitous routes within the innervation layer.     

Retinotopic organization of central optic projections in Rana pipiens

The retinotopic organization of the anuran visual system has been investigated with the method of selective anterograde transport of horseradish peroxidase (HRP) following retinal lesions. The course of optic axons to specific structures was also confirmed by retrograde transport in the optic tract following HRP injections in the tectum and pretectum. As the optic nerve reaches the optic chiasm, the fibers from each of the four retinal quadrants appear as bands with the nasal (n) quadrant entering the chiasmal anterior pole, followed by ventral (v), temporal (t), and dorsal (d) quadrants. The preoptic nucleus is the first structure to be innervated, followed by the suprachiasmatic nucleus; both are innervated directly from fibers in the dorsal part of the optic nerve, which contains fibers from all the retinal quadrants. Each quadrant expands across the dorsoventral extent of the chiasm at the point where it enters. At this level the quadrants are arrayed along the rostrocaudal axis (as they are later in the marginal optic tract) in the sequence n-v-t-d. Optic fibers then spread across the chiasm, the nasal quadrant splits, taking up positions in the rostral and caudal margins of the optic radiation. Following the split in the nasal representation, the optic tract is transformed into topographically arranged sheets in the marginal optic tract. In the other retinorecipient nuclei, the sheet of optic axons is transformed back into the shape of the retinal hemisphere. Topographic maps of this kind display one of two possible orientations: (1) in the tectum and the nucleus lentiformis mesencephali (nLM), the temporal retina is represented in the anterior portion of the nucleus, whereas the nasal quadrant is found in the posterior portion; (2) in the thalamus, the retinotopic map is organized as a mirror-image reversal of that seen in the tectum and nLM (i.e., the nasal pole is anterior, whereas the temporal pole is in the posterior portion of the nucleus). Structures with this type of retinal map include the rostral visual nucleus, the corpus geniculatum, the nucleus of Bellonci, and the posterior thalamic nucleus. A third type of innervation occurs in the nucleus of the basal optic root (nBOR), which is the only mesencephalic visual nucleus not innervated by the marginal optic tract. The basal optic root is formed by the fibers exiting most caudally from the optic chiasm. All the retinal quadrants contribute to the basal optic root, but no evidence of retinotopy was found in nBOR.4+ target nuclei.     

Retinal recipient nuclei in the painted turtle,Chrysemys picta: An autoradiographic and HRP study

Retinofugal pathways in the painted turtle were examined with autoradiographic and HRP methods. The majority of the retinal fibers decussate at the optic chiasm and course caudally to terminate in 12 regions of the diencephalon and mesencephalon. The pars dorsalis of the lateral geniculate nucleus is the densest target in the thalamus. Two nuclei dorsal to pars dorsalis—the dorsal optic and dorsal central nuclei—receive optic input. Three nuclei ventral to pars dorsalis are retinal targets—the ventral geniculate nucleus, nucleus ventrolateralis pars dorsalis, and nucleus ventrolateralis pars ventralis. Contralateral fibers course through the pretectum where they terminate in nucleus geniculatis pretectalis, nucleus lentiformis mesencephali, nucleus posterodorsalis, and the external pretectal nucleus. Retinal fibers also terminate within the superficial zone of the optic tectum. HRP material demonstrates three optic fiber layers—laminae 9, 12, and 14. Optic fibers leave the main optic tract as a distinct accessory tegmental optic pathway and terminate in the basal optic nucleus. Ipsilateral retinal terminals occur in a pars dorsalis and a pars ventralis of the lateral geniculate nucleus, the dorsal optic nucleus, nucleus posterodorsalis, the basal optic nucleus, and in laminae 9 and 12 of the optic tectum. Rostrally, the ipsilateral tectal fibers occupy two zones along the medial and lateral tectal roof; these zones converge caudally and are continuous along the caudal wall of the tectum.     

Optic synapse number but not density is constrained during regeneration onto surgically halved tectum in goldfish: HRP-EM evidence that optic fibers compete for fixed numbers of postsynaptic sites on the tectum

The number of optic synapses in the half tectum of goldfish was counted by using an improved HRP-labeling protocol and a columnar sampling method that spanned the entire optic innervation layer, S-SO-SFGS. It was previously found by using this procedure in intact tectum that the normal number of optic synapses was regenerated by 30 days and maintained thereafter even in the absence of impulse activity. This suggested that the number of synapses in this system was intrinsically fixed. In order to examine whether this limit was imposed by optic fibers or by target cells, optic synapses were counted in surgically halved tecta which received compressed optic projections consisting of regenerating optic fibers from the entire retina. We reasoned that if synapse number is a function of the number of afferents, then there should be twice the normal number of optic synapses per column; on the other hand, if their number is fixed by target, then their number per column should be normal. We found that the number of optic (labeled) synapses was normal in sample columns from fish at 70 days and 160 days after optic nerve crush. Thus, retinal ganglion cells, on average, formed half as many synapses on the half tectum compared to intact tectum, indicating the number of optic synapses was limited by the tectum. The number of nonoptic (unlabeled) synapses was also found to be normal. By contrast, the S-SO-SFGS was found to be 88-103% thicker compared to normal fish, apparently because of a 20-fold increase in the number of optic fibers. As a result, the density of synapses was about half normal in half tecta, and so, in contrast to synapse number, synaptic density is not constrained during regeneration. We infer from these data that optic fibers compete for limited numbers of postsynaptic sites during regeneration and suggest that this competition promotes neural map refinement and the various plasticities described for this projection.