The ground level visual field of each eye of the leopard frog includes the entire ipsilateral 180-deg field and approximately 60 deg into the frontal contralateral field. When one eye is covered with an opaque patch, a frog responds to prey stimuli over the entire field of the other eye. Nevertheless, when one optic nerve is cut, the animal responds to prey in the ipsilateral hemifield of the connected eye, but only responds as far as about 30 deg past the frontal midline. If one optic tract is cut, the animal does not respond to prey past the frontal midline. We hypothesized that the responses past the frontal midline might be mediated by input from contralaterally projecting isthmotectal fibers. These fibers originate in the nucleus isthmi, a posterior midbrain structure. We found that when we placed an opaque patch over one eye and either ablated the ipsilateral nucleus isthmi, or cut crossing isthmotectal fibers in the optic chiasm, or blocked input to nucleus isthmi by ablating the ipsilateral tectal lobe, animals did not respond to prey stimuli past the frontal midline. We found that when we placed an opaque patch over one eye and cut crossing optic fibers in the anterior part of the optic chiasm (sparing crossing isthmotectal fibers), animals responded to prey stimuli in the nasal half of the seeing eye's contralateral frontal field. Our results suggest that contralaterally projecting isthmotectal fibers enable the frog to respond to stimuli past the frontal midline. We suggest a one-dimensional model of how nucleus isthmi influences tectal function.

The retinal topography of the cat's optic tract was determined by means of injections of the enzyme horseradish peroxidase (HRP) into the tract. This analysis was accomplished by the subtraction of all HRP injection sites not labeling a defined retinal area from those injection sites which resulted in ganglion cell labeling (Venn diagram analysis). Using this method, the following correspondences were demonstrated for the ipsilateral and contralateral projections: superior retina represented in medial optic tract; inferior retina in lateral tract; and area centralis in a dorsocentral location (which was part of a larger area representing the visual streak). The temporal raphe was represented in the ipsilateral tract as a band curving from the area centralis region toward the dorsomedial border of the tract. Contralateral fibers from a region superior to the optic disc were found to be displaced with respect to the general retinal representation in the optic tract and this appeared to be related to retinal development. The ratio of contralateral to ipsilateral fibers was determined and found to be nonuniform within the tract.

Injection of HRP into the optic tract of the cat also allowed the axons from labeled retinal ganglion cells to be traced within the retina and optic disc. Axons from ganglion cells lying temporal to the raphe curve around the area centralis enter the optic disc on the lateral and inferior aspects. Ganglion cells lying nasal to the raphe send their axons more directly to enter the optic disc on its superior aspect. A schema is proposed whereby the retina is mapped onto the optic tract.

Introduction

Classical views of the optic chiasm maintain four propositions about the retinofugal pathways: (1) each optic nerve contains a retinotopic representation of its respective retinal surface; (2) this retinotopic map in the nerve is the basis for the subsequent segregation of the decussating from the non-decussating fibers; (3) this retinotopy in the nerve is also the basis for the presence of retinotopy found within the half-retinal maps in the optic tracts; and (4) the half-retinal maps from each optic nerve are brought together within the chiasm to yield a unified, binocularly congruent, map in the optic tract (Brodal, 1969; DukeElder, 1961; Polyak, 1957; Wolff, 1940). The appeal of this classical view is in its simplicity, based on the assumption that the retinofugal pathway should replicate the sensory surface along its course. We now know that each of these four propositions is incorrect, and that the error is not one simply of degree or extent (Guillery, 1982, 1991). Rather, the above description of the visual pathway is fundamentally flawed because it has failed to take into account the constraints under which the pathway develops. We shall first consider the evidence for rejecting the classical view, from recent studies on the organization of the retinofugal pathway in adult animals and on the development of that organization. We shall then describe three transformations in the fiber order which all occur in the chiasmatic region, two of which were only recently recognized, and for which we must account.

Observations from adult organization

The difference in the fiber order in the optic nerve and tract

This study has examined the representation of the dorso-ventral retinal axis in the optic nerve and tract of the ferret, as well as the associated fiber transformations which take place within the chiasmatic region. In one series of experiments, dorsal or ventral retinal lesions were made to induce fiber degeneration along the pathway, from which semi-thin sections were then stained for degenerating myelin. In a second series, implants of the carbocyanine dye, Dil, were made into the caudo-medial or rostro-lateral optic tract in order to label retrogradely the axons as they course through the chiasmatic region. Additional observations were made from the optic pathways of ferrets that had been similarly lesioned or implanted, but employing either a reduced-silver technique to reveal the degenerating axons or horseradish peroxidase as the retrograde label.

The axons arising from the dorsal and ventral retina course in the dorsal and ventral parts of the optic nerve posterior to the eye, but as they continue along the nerve they disperse producing a highly impoverished retinotopy in the prechiasmatic portion of the nerve. As they course through the chiasmatic region, however, they become segregated again: dorsal fibers cross the midline relatively caudally while ventral fibers cross further rostrally, although there is overlap between them. Nearer the threshold of the optic tract, the fibers from dorsal and ventral retina undergo a further and more striking segregation, placing the dorsal fibers caudo-medially and the ventral fibers rostro-laterally within the tract. This re-emergence of retinotopic order implicates a fiber-substrate interaction as being responsible for the axonal reordering, and suggests that fiber pre-ordering in the tract contributes to the formation of the orderly projection of the dorso-ventral retinal axis upon central visual targets.

Brain-derived neurotrophic factor (BDNF) is a preferred ligand for a member of the tropomyosin-related receptor family, trkB. Activation of trkB is implicated in various activity-independent as well as activity-dependent growth processes in many developing and mature neural systems. In the subcortical visual system, where electrical activity has been implicated in normal development, both differential survival, as well as remodeling of axonal arbors, have been suggested to contribute to eye-specific segregation of retinal ganglion cell inputs. Here, we tested whether BDNF is required for eye-specific segregation of visual inputs to the lateral geniculate nucleus and the superior colliculus, and two other major subcortical target fields in mice. We report that eye-specific patterning is normal in two mutants that lack BDNF expression during the segregation period: a germ-line knockout for BDNF, and a conditional mutant in which BDNF expression is absent or greatly reduced in the central nervous system. We conclude that the availability of BDNF is not necessary for eye-specific segregation in subcortical visual nuclei.

Tree shrews (Tupaia belangeri) are small diurnal mammals capable of quick and agile navigation. Electroretinographic and behavioral studies have indicated that tree shrews possess very good temporal vision, but the neuronal mechanisms underlying that temporal vision are not well understood. We used single-unit extracellular recording techniques to characterize the temporal response properties of individual retinal ganglion cell axons recorded from the optic tract. A prominent characteristic of most cells was their sustained or transient nature in responding to the flashing spot. Temporal modulation sensitivity functions were obtained using a Gaussian spot that was temporally modulated at different frequencies (2–60 Hz). Sustained cells respond linearly to contrast. They showed an average peak frequency of 6.9 Hz, a high-frequency cutoff at 31.3 Hz, and low-pass filtering. Transient cells showed nonlinear response to contrast. They had a peak frequency of 19.3 Hz, a high-frequency cutoff at about 47.6 Hz, band-pass filtering, and higher overall sensitivity than sustained cells. The responses of transient cells also showed a phase advance of about 88 deg whereas the phase advance for sustained cells was about 43 deg. Comparison with behavioral temporal modulation sensitivity results suggested that transient retinal ganglion cells may underlie detection for a wide range of temporal frequencies, with sustained ganglion cells possibly mediating detection below 4 Hz. These data suggest that two well-separated temporal channels exist at the retinal ganglion cell level in the tree shrew retina, with the transient channel playing a major role in temporal vision.