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S B Udin

Publications and source records attributed to S B Udin.

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N-methyl-D-aspartate antagonists prevent interaction of binocular maps in Xenopus tectum.

Glutamate receptors appear to play a key role in several forms of experience-dependent modification of both the strength of synapses and synaptic connectivity. In developing Xenopus frogs, the connections made by isthmotectal axons relaying visual input from the eye to the ipsilateral tectum are markedly influenced by the visual activity of contralateral retinotectal axons, and normal binocular visual input is necessary in order for the ipsilateral visuotectal map to come into register with the contralateral map. We have tested whether NMDA receptors play a role in establishment of the topographic matching of binocular maps during development. We have examined the effects of chronic treatment of tectum with either the receptor agonist NMDA or the antagonists APV or CPP applied throughout early postmetamorphic life using subpial implants of drug-impregnated elvax. Both antagonists blocked the matching of the ipsilateral map to the contralateral map, while NMDA permitted such matching. Our data therefore indicate that NMDA receptors are involved in the experience-dependent establishment of matching binocular maps during development.

2-Amino-5-phosphonovalerate↗

Connections between the nucleus isthmi and the tectum in larval and post-metamorphic axolotls.

The nucleus isthmi (NI) is the primary relay for the frog's ipsilateral visuotectal projection. Using electrophysiological methods, ipsilateral visuotectal activity has been recorded in thyroxine-treated, postmetamorphic axolotls but not in larval axolotls. In order to determine whether changes in isthmotectal projections are responsible for this change in electrophysiological responsiveness, we have investigated the connections between the tectum and the NI using horseradish peroxidase. Our results indicate that the axolotl's isthmotectal pathways are strikingly similar to those of the frog NI, and that the NI sends bilateral projections to the tecta in both larval and thyroxine-treated, postmetamorphic axolotls. Thus, the anatomical connections underlying the ipsilateral visuotectal projection are present during larval stages, despite the lack of electrophysiological evidence for the larval ipsilateral visuotectal projection. We hypothesize that thyroxine-induced metamorphosis produces changes in the terminal arborizations of the crossed isthmotectal projection which allow them to be detected by presynaptic electrophysiological techniques.

Ambystoma↗

Frogs use retinal elevation as a cue to distance.

We have investigated the role that retinal elevation plays in a frog's (Rana pipiens) estimate of prey distance. We dissociated retinal elevation from other depth cues by artificially increasing the height of the frogs' eyes above the ground. Frogs then snapped short of their prey in their ventral visual field as if their estimate of distance were determined primarily by the retinal elevation of the image of the prey. The data suggest that the frog assumes its eyes to be about 3 cm above the ground. Other cues modify depth judgements when targets are close to this assumed ground-plane.

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Formation of topographic maps.

The catalogue of data presented here form many systems demonstrates that multiple mechanisms are involved in the formation of topographic maps. We are not yet in a position to explain why a particular mechanism appears to dominate in some situations and not in others. Certain generalizations can be made, however. First, at least some form of chemospecificity can be invoked to help explain connectivity in all of the experiments we have cited. Often, the differential identities of a population of neurons can be reflected in an orderly pattern of axon outgrowth and in the actively maintained preservation of neighbor relations as the axons grow toward their targets; such orderly arrangements are not obligatory, but, where present, they facilitate the speedy establishment of orderly maps when the axons reach their target nuclei. Within a terminal zone, chemospecific cues may dominate and constrain a given axon to terminate in a specific location, but axon-axon interactions commonly supercede chemospecific matching. At least two forms of axon-axon interaction occur, one based on some sort of biochemical properties related to the axon's embryological identity and another based on the axons' electrical activity. Tasks for the future are to identify the cellular bases of each of these mechanisms and to understand the situations in which each is manifested.

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A possible mechanism for binocular depth judgements in anurans.

Two experiments were performed to analyze how anurans (Bufo marinus) use binocular cues to gauge the distance of their prey. In the first, bilateral lesions of the nucleus isthmi eliminated the major source of input from the ipsilateral eye to the tectum. These lesions did not disrupt the animals' ability to use binocular cues to judge distance, suggesting that frogs and toads may not employ binocular disparity-selective cells to assess prey distance. They may instead use a scheme more overtly akin to triangulation, with each tectum providing an output signal encoding the angular position of the prey with respect to the contralateral eye and with distance extracted from the difference between these tectal outputs. In the second experiment, prisms imposed large (13.5 degrees) vertical disparities between the two eyes' images. The toads continued to use binocular cues. The added vertical disparities, like added horizontal ones, caused toads to undershoot their prey. Thus the binocular system must tolerate such vertical disparities and fail to distinguish them from horizontal ones.

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A projection from the mesencephalic tegmentum to the nucleus isthmi in the frogs, Rana pipiens and Acris crepitans.

The nucleus isthmi is a prominent part of the frog's visual system. Each nucleus isthmi receives input from the ipsilateral tectum and sends output to both tecta. Until now, no non-tectal inputs to the nucleus isthmi of amphibians have been demonstrated. Anterograde and retrograde tracing with horseradish peroxidase in Rana pipiens and Acris crepitans now reveal that a diffuse group of cells in the mesencephalic tegmentum projects to the caudal region of the contralateral nucleus isthmi. These cells are primarily within the nucleus anterodorsalis tegmenti. This same group of tegmental cells may also project to the caudal region of the ipsilateral nucleus isthmi. A similar investigation of the brain of another frog, Xenopus laevis, has not revealed any evidence of this tegmento-isthmic projection.

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The development of the nucleus isthmi in Xenopus laevis. I. Cell genesis and the formation of connections with the tectum.

The nucleus isthmi (NI) of the amphibian relays visual input from one tectum to the other tectum and thus brings a visual map from the eye to the ipsilateral tectum. This isthmotectal visual map develops slowly; it is first detected electrophysiologically at stages 60-62, the age at which the eyes begin their dorsalward migration and the region of binocular overlap beings to increase in extent. During this critical period of life, normal binocular visual input is required for establishment of normal topographic isthmotectal projections. In this study, we have used anatomical methods to trace cell birth, cell death, and formation of connections by the nucleus isthmi during the critical period. Tritiated thymidine labelling demonstrates that cells in the nucleus isthmi are generated throughout most of tadpole life (stages 29-62). Most cells conform to an orderly ventrodorsal gradient starting from stage 29 and extending to stages 56; later cells are inserted at apparently random locations in the nucleus. We have re-examined the hypothesis of Tay and Straznicky ('80) that the order of cell genesis in the NI and tectum could help establish proper isthmotectal connections, and we find that a timing mechanisms does not explain the two-dimensional topography of the isthmotectal map but that timing may aid in proper mediolateral positioning of isthmotectal axons at the points where they first enter the tectum. Horseradish peroxidase labelling was used to investigate whether anatomical projections from tectum to NI and from NI to tectum are present prior to the onset of eye migration. The results show that there are tectoisthmotectal projections by stage 52. Moreover, isthmotectal axons grow into as yet monocular tectal regions prior to the onset of eye migration. At stage 60, when binocular overlap begins, isthmotectal axons are visible throughout the tectum but are densely branched only at the rostral tectal margin, the location where they are predicted to occur on the basis of electrophysiological maps.

Acetylcholinesterase↗

The role of visual experience in the formation of binocular projections in frogs.

Many parts of the visual system contain topographic maps of the visual field. In such structures, the binocular portion of the visual field is generally represented by overlapping, matching projections relayed from the two eyes. One of the developmental factors which helps to bring the maps from the two eyes into register is visual input. The role of visual input is especially dramatic in the frog, Xenopus laevis. In tadpoles of this species, the eyes initially face laterally and have essentially no binocular overlap. At metamorphosis, the eyes begin to move rostrodorsally; eventually, their visual fields have a 170 degree region of binocular overlap. Despite this major change in binocular overlap, the maps from the ipsilateral and contralateral eyes to the optic tectum normally remain in register throughout development. This coordination of the two projections is disrupted by visual deprivation. In dark-reared Xenopus, the contralateral projection is nearly normal but the ipsilateral map is highly disorganized. The impact of visual input on the ipsilateral map also is shown by the effect of early rotation of one eye. Examination of the tectal lobe contralateral to the rotated eye reveals that both the contralateral and the ipsilateral maps to that tectum are rotated, even though the ipsilateral map originates from the normal eye. Thus, the ipsilateral map has changed orientation to remain in register with the contralateral map. Similarly, the two maps on the other tectal lobe are in register; in this case, both projections are normally oriented even though the ipsilateral map is from the rotated eye. The discovery that the ipsilateral eye's map reaches the tectum indirectly, via a relay in the nucleus isthmi, has made it possible to study the anatomical changes underlying visually dependent plasticity. Retrograde and anterograde tracing with horseradish peroxidase have shown that eye rotation causes isthmotectal axons to follow abnormal trajectories. An axon's route first goes toward the tectal site where it normally would arborize but then changes direction to reach a new tectal site. Such rearrangements bring the isthmotectal axons into proximity with retinotectal axons which have the same receptive fields. Anterograde horseradish peroxidase filling has also been used to study the trajectories and arborizations of developing isthmotectal axons. The results show that the axons enter the tectum before the onset of eye migration but do not begin to branch profusely until eye movement begins to create a zone of binocular space.(ABSTRACT TRUNCATED AT 400 WORDS)

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Intertectal neuronal plasticity in Xenopus laevis: persistence despite catecholamine depletion.

In normal Xenopus, the tectum receives a direct projection from the contralateral retina and an indirect projection, via the intertectal system, from the ipsilateral eye. The two maps of binocular visual space, at each tectum, are in register. If one eye is rotated during larval development, the ipsilateral visuotectal projection compensates by changing its orientation. Rearrangement of the intertectal system brings the ipsilateral map back into register with the contralateral map. We sought to determine whether this intertectal plasticity required normal levels of brain monoamines. Animals received an eye rotation between stages 55-63 of larval life and were then placed in one of 3 groups. A first control group received no further treatment. A second control group was given intraventricular injections of ascorbate vehicle. The experimental group was given intraventricular injections of 6-hydroxydopamine in ascorbate vehicle. Two to 3 months after metamorphosis, visuotectal projections were mapped electrophysiologically and the brains were assayed for monoamines. Intertectal plasticity occurred in all 3 groups of animals, including animals in which brain catecholamine levels were severely reduced. We conclude that normal levels of brain catecholamines are not required for this form of neural plasticity.

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Abnormal visual input leads to development of abnormal axon trajectories in frogs.

Throughout the normal vertebrate brain, visual maps form the left and right eyes overlap and are in register with one another. Visual input has a major role in the development of the pathways which mediate these binocular projections. A dramatic example of the developmental role of sensory input occurs in the isthmo-tectal projection, which is part of the polysynaptic relay from the eye to the ipsilateral tectum of the frog, Xenopus laevis. If one eye is rotated when the animal is still a tadpole, the isthmic axons respond by changing the topography of their terminations in the tectum; for example, a given isthmo-tectal axon which normally would connect with medial tectum can be induced to terminate in lateral tectum. Such rearrangements bring the ipsilateral visual map into register with the contralateral retinotectal map, even though one eye has been rotated. Indirect evidence has suggested that after early eye rotation, isthmo-tectal axons do not grow directly to their new tectal targets but instead reach those targets by routes which pass through their normal termination zones. Here I have used anterograde horseradish peroxidase labelling of isthmo-tectal fibres to show the trajectories of such axons and to compare them with the routes which axons take when allowed to develop normally. Tracings of individual axons in flat-mounted, unsectioned tecta show that most axons in normal Xenopus follow fairly straight paths in the tectum. In contrast, early eye rotation causes many isthmo-tectal axons to follow crooked, circuitous pathways before they terminate.

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Expansion and retinotopic order in the goldfish retinotectal map after large retinal lesions.

A peripheral annulus of neural retina and pigment epithelium was removed from the eye of adult goldfish. After survival times from 148 to 560 days, the retinotectal projection from the remaining central fragment was mapped. In most cases, the map was orderly and had expanded to fill the entire contralateral tectum, but when less than 10-15% of the original retina remained intact, the projection failed to fill the entire available tectal space and was abnormally disordered.

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Plasticity in a central nervous pathway in xenopus: anatomical changes in the isthmotectal projection after larval eye rotation.

In this paper we present evidence that early eye rotation in Xenopus leads to anatomical rearrangements in a portion of the binocular visual system. In the past, electrophysiological mapping had shown that the topography of the ipsilateral visuotectal projection is changed by such eye rotation and that this change requires visual experience. However, knowledge of the anatomical basis for this electrophysiological change was lacking. The identification of the nucleus isthmi as a link in the projection has allowed us to study the topography of the ipsilateral system by use of horseradish peroxidase. We present data showing that early eye rotation alters the topography of the crossed isthmotectal projection. These results demonstrate that the orientation of a topographically organized projection can be changed by procedures which do not involve direct manipulation of the source, pathway, or target of the projection.

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Removal of retinal input fails to elicit isthmo-tectal sprouting in the frog.

The retina and nucleus isthmi both project in laminar fashion to the superficial layers of the frog's tectum. In order to determine whether isthmo-tectal axons show collateral sprouting after retinotectal input is removed, we injected [3H] proline into the nucleus isthmi and measured the volume of the crossed isthmo-tectal projection. We found no evidence of collateral sprouting.

Afferent Pathways↗

Compressed retinotectal projection in hamsters: fewer ganglion cells project to tectum after neonatal tectal lesions.

After partial ablation of the superior colliculus (tectum) in neonatal hamsters, the whole extent of the visual field comes to be represented in a compressed map on the remaining tectal fragment. However, the total volume of tectal tissue in which retinotectal fibers arborize is less than normal. These observations suggests that the retinal ganglion cells which arborize in this reduced volume might arise throughout the whole extent of the retina but be fewer in number than normal. Alternatively, the ganglion cells which project to the tectum might be normal in number but reduced in terminal arbor size. To distinguish between these possibilities, we have used tectal injections of horseradish peroxidase to label retinal ganglion cells which project to the tectum. The numbers of labelled cells per mm2 of retina were counted in selected regions. In hamsters with small lesions, which left 80--85% of the tectum intact, the density of labelled retinal ganglion cells was normal. However, in hamsters with larger lesions, the density of labelled cells was significantly lower than normal.

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Topographic projections between the nucleus isthmi and the tectum of the frog Rana pipiens.

The connections between the nucleus isthmi and the tectum in the frog have been determined by several anatomical techniques: iontophoresis of horseradish peroxidase into the tectum, iontophoresis of 3H-porline into the nucleus isthmi and the tectum, and Fink-Heimer degeneration staining after lesions of the nucleus isthmi. The results show that the nucleus isthmi projects bilaterally to the tectal lobes. The ipsilateral isthmio-tectal fibers are distributed in the superficial layers of the tectum, coincident with the retionotectal terminals. The contralateral isthmio-tectal fibers travel anteriorly adjacent to the lateral optic tract and cross the midline in the supraoptic ventral decussation, where they turn dorsally and caudally; upon reaching the tectum, the fibers end in two discrete layers, layers 8 and A of Potter. The tectum projects to the ipsilateral nucleus isthmi and there is a reciprocal topographic relationship between the two structures. Thus, a retino-tecto-isthmio-tectal route exists which may contribute to the indirect ipsilateral retinotectal projection which is observed electrophysiologically. The connections between the nucleus isthmi and the tectum in the frog are strinkingly similar to the connections between the parabigeminal nucleus and the superior colliculus of mammals.

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Rearrangements of the retinotectal projection in Rana pipiens after unilateral caudal half-tectum ablation.

When the caudal half-tectum is ablated and the optic nerve is cut in adult Rana pipiens, recordings from optic nerve terminals show that the entire visual field comes to be represented in retinotopic order on the remaining rostral half-tectum (compression). In contrast, if the caudal half-tectum is ablated but the optic nerve is left intact, no compression of the retinotectal projection results. Instead, the terminals displaced from the ablated tissue form a permanently disorganized projection to the rostral half-tectum superimposed on the unaltered representation of those parts of the retina which normally project there. The receptive field locations of tectal neurons in both groups of animals show that the altered retinotectal projections make functional synapses. This conclusion is further supported by behavioral data, which show that the accuracy of prey-catching movements is altered in both groups of frogs. The anomalous retinotectal maps can be interpreted by means of a model which includes polarity cues derived from the tectum and ordering cues derived from repelling interactions among fiber terminals.

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Differential intertectal delay between Rana pipiens and Xenopus laevis: implications for species-specific visual plasticity.

In the frog Xenopus laevis, the isthmotectal projection, which relays input from the ipsilateral eye, exhibits anatomical reorganization following surgical eye rotation performed during tadpole stages while the isthmotectal projection in the frog Rana pipiens fails to show reorganization. This plasticity has been shown to be dependent upon activation of the N-methyl-D-aspartate (NMDA) receptor located on tectal cell dendrites. The reorganization process in Xenopus is hypothesized to employ a Hebbian mechanism requiring correlated firing of ipsilateral and contralateral inputs to a given tectal cell; when an ipsilateral axon synapses onto a tectal cell that receives input from a contralateral axon with a matching receptive-field location, the correlation in activity triggers stabilization of the ipsilateral synapse. However, in neither Xenopus nor Rana do ipsilateral and contralateral inputs begin to fire simultaneously in response to a given visual stimulus; the ipsilateral input is delayed because it reaches the tectum indirectly, through a polysynaptic relay via the opposite tectum and nucleus isthmi. The objective of this experiment was to test whether there is a significant difference in this intertectal delay between Xenopus laevis and Rana pipiens in order to determine whether intertectal delay could be a contributing factor in this species-specific ability to exhibit visual plasticity. We have found that intertectal delay is 26.16 ms longer in Rana pipiens (36.53 ms) than in Xenopus laevis (10.37 ms).

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