A degeneration study of the course and extent of the pyramidal tract of the opossum.
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Retinal projections were examined in the native cat, Dasyurus viverrinus using Fink-Heimer material and autoradiography. We found six regions in the brain which receive retinal projections. These are (1) the dorsal lateral geniculate nucleus (2) the ventral lateral geniculate nucleus (3) the lateral posterior nucleus (4) the pretectum (5) the superior colliculus, and (6) the accessory optic system. We did not examine the hypothalamus. The accessory optic system and the lateral posterior nucleus receive a contralateral retinal projection only and the other four regions receive a bilateral retinal projection. There is extensive binocular overlap in the dorsal lateral geniculate nucleus. On the side contralateral to an eye injection of 3H leucine our autoradiographs show four contralateral layers which fill most of the nucleus. Three of these layers, 3, 4 and 5, also receive input from the opsilateral eye. Layer 1 which lies adjacent to the optic tract receives only contralateral retinal input. Layer 2 receives a direct retinal input only from the ipsilateral eye. The ipsilateral projection to the dorsal lateral geniculate nucleus forms a fairly continuous patch which is not divided into separate layers. The ipsilateral retinal input is located in the dorsal part of the lateral geniculate nucleus. The ventral quarter of the nucleus only receives a contralateral retinal input and therefore represents the monocular part of the visual field.
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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.
Maps of ganglion-cell distribution have been constructed from whole-mounted retinas of five Australian marsupial species. The pademelon wallaby (Thylogale billiardieri), the scrub wallaby or tammar (Macropus eugenii), and the carnivorous Tasmanian devil (Sarcophilus harissi) have both a visual streak and an area centralis. The retina of the brown bandicoot (Isoodon obesulus) also shows both these features but they are less prominent than in the former three species, whereas the burrow-dwelling, hairy-nosed wombat (Lasiorhinus latifrons) possesses a well-developed visual streak but seems to lack an area centralis. A study of ganglion-cell sizes comparing nasal and temporal retina, the visual streak, and/or the area centralis was undertaken in each species. Results show that as in the cat, small ganglion cells tend to concentrate in the visual streak. However, the temporal-nasal differences in cell sizes described in the cat (Stone et al., '80) could be detected only in those marsupials in which an area centralis was clearly recognizable.
The distribution, morphology, size, and number of cells in the retinal ganglion layer of the marsupial Setonix brachyurus, "quokka," was studied from 25 days postnatal to adulthood using Nissl-stained wholemounts. The total cell population was evenly distributed up to 50 days, but by 75 days highest densities were generally observed in a broad band extending across the nasotemporal axis. At 87 days, a temporally situated area centralis was seen for the first time. This was embedded in a horizontally aligned visual streak, the nasal arm of which contained areas of high density. By 106 days, densities in the area centralis had stabilized while peripheral values were higher than adult levels even at 180 days. In the adult, the area centralis was surrounded by a weak visual streak. Retinal area increased steadily during development to reach 168 mm2 at 180 days, the adult range being 225-250 mm2. All cells in the ganglion layer appeared undifferentiated and rounded at 33 days with soma diameters of 3-6 micrometers; by 70 days diameters had increased to 4-12 micrometers and some cells had axon hillocks containing Nissl substance. From 87 days we distinguished ganglion cells, which constituted 54-63% of the total. These were identified by deeply stained Nissl substance and had diameters of 7-18 micrometers, compared to 7-23 micrometers at 143 days and 7-24 micrometers in the adult; the remaining cells, termed glia/interneurons, were 5-8 micrometers throughout. Only ganglion cells were organized into an area centralis and visual streak. Glia/interneurons were evenly distributed except at the extreme periphery, where their density increased. In sectioned material, the ganglion layer was distinct from 25 days while the neuroblastic layer separated only between 48 and 85 days. From 25 to 250 days the total number of cells in the ganglion layer remained similar to the adult range of 336,000-393,000. At both 87 days and in adults optic axon counts fell between 180,000 and 224,000, close to ganglionic cell estimates. At 25 and 34 days, respectively, optic axon numbers were 75,000 and 172,000. Myelination was absent at 25 and 34 days, 3% at 87 days, and almost 100% in adults. Mechanisms are discussed whereby the area centralis and visual streak may evolve from an even distribution of cells while their number remains constant; migration is considered likely to be important.
The anatomy of the dorsal lateral geniculate nucleus (LGd) is described in five macropodid species, including two rat kangaroos (bettong and potoroo), two wallabies (pademelon and tammar), and the large grey kangaroo. The distribution of retinal terminals in the LGd was examined following intraocular injections of tritiated amino acids. There are considerable differences in both LGd cytoarchitecture and the patterns of retinal terminations among the five species. Cytoarchitecture in the bettong LGd is relatively simple, displaying a minimal regional differentiation. In contrast, the potoroo LGd is quite complex and displays several well-defined cell laminae, each of which is associated with input from a single eye. Both rat kangaroos display the same basic pattern of retinal termination with three bands of terminals from the contralateral eye and four from the ipsilateral eye. The bands are less sharply defined in the bettong, in which terminals from each eye overlap to a greater extent than is seen in the potoroo. The wallabies and kangaroos display a more complex LGd architecture and patterning of retinal terminal bands. Bilateral retinal projections within the same LGd lamina are unusual in these large macropodids. The number of terminal bands reaches ten in the grey kangaroo--four from the contralateral eye and six from the ipsilateral eye. The pademelon LGd is unusual in that it shows intraspecies variation with some animals displaying five ipsilateral terminal bands and others only four. The results are discussed in comparison with the patterns of LGd organisation observed in other mammalian lines, placental and marsupial. We conclude that LGd lamination and the segregation of retinal inputs to the LGd in marsupials are likely to be the result of evolutionary factors which differ from those which have produced ocular segregation and complex lamination in several lines of placental mammals.
The time course and pattern of retinal innervation of primary visual areas was traced in pouch-young wallabies. Tritiated proline was injected into one eye of animals ranging in age from 1 to 72 days after birth. These results are compared to the 11 primary visual areas found in the adult wallaby, seven of which receive binocular input while four are monocular. At birth retinal ganglion cell axons have not reached any visual areas. Two to 4 days after birth, all of the axons are crossing to the contralateral optic tract. Nine to 12 days after birth axons begin to invade the contralateral lateral geniculate nucleus, the superior colliculus, and the medial terminal nucleus. Twenty to 21 days after birth, ipsilateral axons invade the lateral geniculate nucleus and superior colliculus. The contralateral projection precedes the ipsilateral projection in all binocular visual areas. By 25 days, ipsilateral and contralateral afferents share common territory in the lateral geniculate nucleus; however, afferents from each eye are initially concentrated in appropriate areas. Between 52 and 72 days, afferents to the dorsal lateral geniculate nucleus are gradually segregated into nine terminal bands. Four are contralateral while five are ipsilateral. By 72 days, the ipsilateral component to the superior colliculus is clustered beneath the contralateral projection a deeper layer. Projections to four monocular visual areas--lateral posterior nucleus, dorsal terminal nucleus, lateral terminal nucleus, and nucleus of the optic tract--are established later than binocular visual areas, except the suprachiasmatic nucleus. The suprachiasmatic nucleus is the last to be bilaterally innervated even though it is situated closest to the optic chiasm. At the light microscope level a mature pattern of visual development is emerging by 72 days, although the eyes do not open until 140 days.