Biomedical subjects

L D Beazley

Publications and source records attributed to L D Beazley.

At least 55 records · Page 3Linked to original sources

PubMed1995

Substance P, bombesin, and leucine-enkephalin immunoreactivities are restored in the frog tectum after optic nerve regeneration.

Extensive regeneration of the optic nerve takes place in adult Amphibia. In this study, we have determined whether one aspect of retinotectal organisation, namely immunoreactive laminae in the retinorecipient layers of the optic tectum, is restored after optic nerve regeneration. To do so, the distributions of substance-P, bombesin, and leucine-enkephalin immunoreactivities were examined in the optic tectum of the frog Litoria (Hyla) moorei. Results of a normal series were compared with those at intervals up to 84 days and at 196 days after either unilateral deafferentation or optic nerve crush. In the normal series, distinct neuropeptide immunoreactive laminae were located within the retinorecipient tectal layers. There were two major laminae with substance-P, two with bombesin, and one with leucine-enkephalin immunoreactivities. Additional faint laminae of both substance-P and bombesin immunoreactivity were present in the tectal region that receives input from the visual streak. In addition, labelling of cell bodies and dendrites was seen elsewhere in the tectum. All except one immunoreactive lamina changed after deafferentation. The deeper of those with substance-P immunoreactivity, along with both bombesin laminae, were eventually lost; the lamina with leucine-enkephalin immunoreactivity was halved in intensity. We assume that these laminae are wholely or, in the case of the leucine-enkephalin lamina, partially associated with primary optic input. By contrast, the more superficial lamina with substance-P immunoreactivity remained unchanged and is presumably not directly related to visual input. During nerve regeneration, the intensity of all laminae associated with optic input initially fell as in the deafferentation series but, in the long term, recovered to approximately 80% of normal intensities. We conclude that ganglion cells associated with each of the immunoreactivities tested had successfully regenerated. The reduced intensity of immunoreactivities after regeneration is due presumably in part to the cell loss from the ganglion cell population. Furthermore, we discuss the findings of similar studies for Rana pipiens (Kuljis and Karten [1983] J. Comp. Neurol. 217:239-251 and [1985] 240:1-15) in light of the present findings. We argue that some of the previous observations can be reinterpreted to indicate that regeneration was not limited to ganglion cells associated with substance-P immunoreactivity as first thought.

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PubMed1995

Wiring up the visual system.

1. In this review we describe some of our recent studies on the developing marsupial visual pathway. The description focuses on retinal ganglion cells, considering the formation of their dendritic trees, the outgrowth of axons and the formation of connections within the brain. 2. Both dendritic trees and outgrowing axons undergo a period of exuberance, followed by one of refinement. The dendritic tree transiently develops a more complex branching pattern than is found in adults. Short side branches, referred to as spines, are a feature of immature dendrites and, to a lesser extent, of axons. These structures are mostly lost as development proceeds. However, they are retained on the dendritic trees of small-field ganglion cells and, for a proportion of axons, on that part within the nerve fibre layer of the retina. Although most axons navigate fairly direct routes towards their targets, a minority follow inappropriate courses, such as doubling back towards the eye or entering the opposite optic nerve at the chiasm. As such errant axons are not seen in the adult, we assume that their parent cell bodies die during development. 3. Throughout development, optic axons are arranged in an approximate retinotopic order along the length of the visual pathway; as a result, axons approach the visual centres aligned to form, at least, a crude retinotopic map. Axons from dorsal and ventral retina exchange locations along the optic nerve and in this way correct for the inversion of the image brought about by the lens.(ABSTRACT TRUNCATED AT 250 WORDS)

Adult↗

PubMed1995

Development of the visual cortex in a wallaby--phylogenetic implications.

The visual cortex of one of the smallest macropod marsupials, a wallaby, the quokka Setonix brachyurus, was examined at maturity and during development from postnatal day 1 to 150 in Nissl-stained or Golgi-stained sections. Injections of horseradish peroxidase into the primary visual centres in adults identified cortical neurons projecting to these regions. The pattern of cell generation was determined by tritiated thymidine/autoradiography. The adult visual cortex was composed of the usual six layers of cells with dendritic morphologies similar to those seen in eutherian (placental) mammals. The margin of Layer 6 overlapped slightly with the white matter, and pyramidal cells projecting to subcortical regions were located at all depths of Layer 6. Cortical development was similar to that of eutherian mammals, with the formation of transient cellular layers below the developing cortical plate. Autoradiographic studies confirmed that the cortical plate was composed of two layers: a compact zone of densely packed cells located pial to a striated layer containing more loosely packed cells. Postnatal cell generation of Layers 6-2 took place from postnatal day 3 to day 85, with Layer 4 neurons, the main target of incoming visual axons, being generated around postnatal day 40. Cells located at maturity in the white matter, mostly of glial morphology, were generated from postnatal day 40. A subplate zone containing early differentiating cells was not evident, and postnatal 3H-thymidine labelling did not identify a population of early generated neurons below the cortical plate. In the tammar wallaby Macropus eugenii, the majority of target neurons for incoming geniculate axons are generated about 20 days after the first axons enter directly into the cortical plate without a waiting period in a subplate. Geniculate axons in the wallabies have a relatively longer 'wait' for their target neurons than do those in cats or monkeys. Therefore, if geniculate axons in marsupials make temporary synapses, while they wait for their target neurons to be generated, their temporary connections must, unlike those in cats or monkeys, be with cells already in the cortical plate.

Aging↗

PubMed1994

The retinal ganglion cell layer and optic nerve in a marsupial, the honey possum (Tarsipes rostratus).

We examined the retina and optic nerve of one of the smallest mammals, the honey possum (Tarsipes rostratus), a marsupial. Ganglion and displaced amacrine cells were distinguished by morphological criteria in retinal wholemounts stained with cresyl violet. There was a mean of 90,000 ganglion cells which were distributed as an area centralis in the mid-temporal retina with a peak density of 9,000/1 mm2 falling in a concentric shallow gradient to 2,000/1 mm2 at the periphery. The unimodal and positively skewed spectra of soma diameters gave no indication of ganglion cell classes. Displaced amacrine cells numbered only 15,000, being outnumbered 6:1 by ganglion cells. This value represents by far the lowest percentage of displaced amacrine cells yet reported for a mammalian retina. The displaced amacrine population was evenly distributed. Counts of optic axons from electron micrographs were within 1-5% of the ganglion cell estimates. In the retrobulbar portion of the nerve, a substantial proportion of axons (26-43%) were unmyelinated, and many of these were concentrated within the lateral quadrant. This specialisation may represent the outflow of the area centralis. At the level of the foramen, the percentage of unmyelinated axons fell to 4-6%, and axons were more uniform in their distribution. Axon diameters were lower for the unmyelinated than the myelinated populations at both levels. As for ganglion cells, however, the spectra of axon diameters were unimodal and positively skewed for both unmyelinated and myelinated axons.

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PubMed1993

Transient neovascularisation of the frog retina during optic nerve regeneration.

In conditions such as diabetic retinopathy, degenerative events in the retina are associated with neovascularisation. It is well established that a proportion of retinal ganglion cells die during optic nerve regeneration in the frog. The present study has determined whether neovascularisation takes place during this regenerative process. To do so, the pattern of blood vessels overlying the retinal ganglion cell layer was analysed in the frog Litoria (Hyla) moorei. We examined normal animals and those undergoing optic nerve regeneration following nerve crush. Blood vessels were visualised by perfusion with Indian ink and retinae were prepared as wholeamounts. In normal animals, the vascular tree was found to lie superficial to the nerve fibre layer and was more complex in regions overlying the area centralis and visual streak. After nerve crush, abnormal blood vessels transiently formed between the existing branches of the vascular tree. The new vessels were concentrated as an annulus centred on the optic nerve head and over the area centralis in midtemporal retina. The neovascularisation became most extensive between 6 and 10 weeks postcrush and disappeared by 12 weeks. The spatiotemporal sequence of neovascularisation suggests that it is triggered by accumulations of degenerating material formed as a proportion of the ganglion cells die during optic nerve regeneration.

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PubMed1992

Biphasic retinal neurogenesis in the brush-tailed possum, Trichosurus vulpecula: further evidence for the mechanisms involved in formation of ganglion cell density gradients.

We investigated cell generation in the retina of the brush-tailed possum (Trichosurus vulpecula) by using tritiated (3H)-thymidine labelling of newly generated cells. Animals aged between postnatal day (P) 5 and 85 each received a single injection of 3H-thymidine. Following autoradiographic processing, maps of labelled cells were constructed from retinal sections. Retinal cell generation takes place in two phases, the first is concluding in the retinal periphery at P53 as the second is seen to commence in midtemporal retina. In the first phase, cells in central retina are generated earlier than those in peripheral regions. In the second phase, cells complete their final division in midtemporal retina first and in the periphery last. Cells generated in the first phase comprise virtually all cells in the ganglion cell layer, amacrine cells, horizontal cells, and cones. Ganglion cells are produced at a slightly earlier stage than displaced amacrine cells, horizontal cells, or cones. Amacrine cells in the inner nuclear layer are the final cells produced in the first phase. When ganglion cells and amacrine cells are pooled, their combined rate of production matches that of the other cell types. These data indicate that the ratio of displaced amacrine cells: horizontal cells: cones: combined ganglion cells and amacrine cells does not change throughout development. However, the ratio of ganglion cells:macrines changes steadily as development proceeds to favour amacrine cells. In the second phase, sparse numbers of nonganglion cells in the ganglion cell layer and large numbers of bipolar and Müller cells are produced along with all rods. The two phases in the possum are similar to those seen in the wallaby, the quokka. However, fewer cells are added in central retina in the possum than in the quokka and cell addition continues for a more extended period in the periphery in the possum. We suggest that this difference in cell addition could account for the development of a more pronounced visual streak of retinal ganglion cells in the possum than in the quokka. A comparison of possum retinal cell generation with that of other marsupials adds support for the "homochrony theory."

Aging↗

PubMed1992

Displaced retinal ganglion cells in normal frogs and those with regenerated optic nerves.

We have analysed the number and spatial distribution of displaced retinal ganglion cells in the frog Litoria (Hyla) moorei. A series of normal animals was compared with one in which the optic nerve was crushed and allowed to regenerate. Ganglion cells were labelled with horseradish peroxidase (HRP) applied to the optic nerve, and retinae were examined as sections or whole mounts. We analysed separately ganglion cells with somata displaced to the inner nuclear (Dogiel cells, DGCs) and to the inner plexiform layer (IPLGCs). These findings were related to data for the orthotopic ganglion cells (OGCs). The mean number of DGCs in the normal series was 2,550 (+/- 281) and fell to 1,630 (+/- 321) after regeneration, representing a mean loss of 36%. This reduction was not significantly different from the mean loss of 43% from the OGC population in which mean values fell from 474,700 (+/- 47,136) to 268,700 (+/- 54,395). In both the normal and the regenerate series, DGCs were estimated to represent means of only 0.6% of the OGC population. Densities of DGCs were highest in the naso-ventral and temporo-dorsal peripheries; densities of both DGCs and OGCs were lower after optic nerve regeneration. We conclude that the factors which affect ganglion cell death during optic nerve regeneration, do so to similar extents amongst the DGC and the OGC populations. The IPLGCs were very rare in normal animals with a mean of 420 (+/- 95). However, their numbers increased after regeneration to a mean of 3,350 (+/- 690), estimated to be 1.2% of the OGC population. These cells normally favoured peripheral retina but became pan-retinal after regeneration.(ABSTRACT TRUNCATED AT 250 WORDS)

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PubMed1992

Disconnected optic axons persist in the visual pathway during regeneration of the retino-tectal projection in the frog.

In this study, we crushed one optic nerve in the frog Litoria (Hyla) moorei and at intervals thereafter anterogradely labelled optic axons with horseradish peroxidase (HRP). For one series, HRP was applied between the eye and the crush site and in a second series between the crush site and the chiasm. A tectal projection of regenerating axons was seen in both series but, in addition, up to 12 weeks post-crush, the second series displayed an additional projection. Its appearance matched that of the disconnected, but persisting, optic axon terminals which are found after enucleation or optic nerve ligation. We conclude that, in the frog, many disconnected optic axons persist throughout the period of optic nerve regeneration and of restoration of an orderly retino-tectal map.

Animals↗

PubMed1992

A breakdown of the blood-brain barrier is associated with optic nerve regeneration in the frog.

We have examined the integrity of the blood-brain barrier during optic nerve regeneration in the frog Litoria (Hyla) moorei using rhodamine B-labeled bovine serum albumin (RBA). A transient localized breakdown of the blood-brain barrier was observed between 1 and 5 weeks after extracranial optic nerve crush. The zone of breakdown progressed along the experimental optic nerve, ascended the opposite optic tract, and swept rostro-caudally across the tectum contralateral to the crushed nerve. By 7 weeks, the blood-brain barrier was once again intact along the length of the optic pathway. In a concurrent series of frogs, regenerating optic axons were visualized by anterograde transport of horseradish peroxidase (HRP). At each stage examined, the region reached by the front of regenerating axons corresponded to that in which the blood-brain barrier had been shown to break down. In contrast to the results after nerve crush, the blood-brain barrier remained intact along the length of the optic pathway following optic nerve ligation to prevent regeneration. We conclude that the breakdown of the blood-brain barrier which occurs during optic nerve regeneration in the frog is triggered by the regenerating axons.

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PubMed1990

Retinal ganglion cell death during regeneration of the frog optic nerve is not accompanied by appreciable cell loss from the inner nuclear layer.

We estimated cell numbers in the ganglion cell and inner nuclear layers of adult frog (Hyla moorei) retinae, examining normal animals and those with regenerated optic nerves. Analysis of sections stained with cresyl violet showed that cell numbers in a nasotemporal strip, which included the area centralis and visual streak, were comparable between sides for both these cellular layers in normal animals. In line with our previous observations, after optic nerve regeneration cell numbers in the ganglion cell layer had fallen by 35-43% compared to the unoperated sides. By contrast cell numbers remained similar for the inner nuclear layers on the two sides. We conclude that retrograde transneuronal degeneration had not taken place in the inner nuclear layer in response to ganglion cell death.

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PubMed1990

Development of topographic connections between the isthmic nuclei and optic tecta in the frog Limnodynastes dorsalis.

In the frog Limnodynastes dorsalis, the pattern of topographic connections between the isthmic nuclei and optic tecta was determined by anterograde and retrograde transport of horseradish peroxidase from localised tectal regions. In both larvae and adults, reciprocal mapping of the uncrossed isthmo-tectal and tecto-isthmal projections was evidenced by the juxtaposition of labelled tecto-isthmal terminations with labelled cells in the cortex and medulla of the ipsilateral isthmic nucleus. The crossed isthmo-tectal projection was revealed by labelled cells in the cortex and medulla of the nucleus contralateral to the injection. In adults, rostral tectal areas projected to rostral and ventral regions of the ipsilateral isthmic nucleus. Following more caudal tectal injections, labelled cells were found in progressively more dorsal locations within the nucleus. Labelled cells in the contralateral nucleus were found in the rim cortex abutting a neuropil and in medullary cells adjacent to this region. Connections between ventral isthmic regions and most rostral tectum and between dorsomedial nucleus and caudomedial tectum were similar in both nuclei. However, for isthmic areas projecting to rostromedial and mid-tectum, the location of labelled cells in the contralateral nucleus was inverted with respect to the ipsilateral nucleus. This inversion would allow both nuclei to project to visually corresponding regions of each tectum. During larval stages the basic adult topography was established despite the continued neurogenesis of both isthmic nuclei and optic tecta. In late larval stages a rim neuropil appeared adjacent to the cortical region in the isthmic nuclei where labelled cells of the crossed isthmotectal projection were found. Prior to this stage labelled cells abutted labelled medullary cells. The appearance of this neuropil was approximately temporally correlated with the onset of electrophysiologically detectable responses in the ipsilateral visuotectal projection. Formation of the rim neuropil may relate to maturation of the tecto-isthmo-tectal connections which underlie this visual projection.

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PubMed1990

Retinofugal projections in a marsupial, Tarsipes rostratus (honey possum).

We examined the mature retinal projections of the honey possum, Tarsipes rostratus, an Australian, diprotodont marsupial, using uni-ocular injections of horseradish peroxidase and tetramethylbenzidine processing. The suprachiasmatic nucleus, medial, lateral and dorsal terminal nuclei of the accessory optic tract, pretectal nuclei and superior colliculus received bilateral retinal input. Contralateral input only was observed in the lateral posterior nucleus. The pattern of input to these regions was essentially similar to that seen in other marsupials. Cyto-architectural examination of the dorsal lateral geniculate nucleus (dLGN) indicated that 5 laminae were present in the alpha-segment, but the beta-segment appeared to lack lamination. Input to the dLGN was bilateral, overlapping considerably, and was organised in a laminar fashion with 7 and 3 terminal bands in the alpha- and beta-segments, respectively. The monocular segment accounted for 12% of the total volume of the dLGN. In the alpha-segment, 2 terminal bands each received exclusively contralateral or ipsilateral input and 3 bilateral input. In the beta-segment, 2 terminal bands received bilateral and 1 contralateral input. The volumes of the nucleus receiving contralateral and ipsilateral input were 77 and 54% of the total, respectively. A marked overlap of input from the two eyes is an unusual feature for a diprotodont marsupial and has previously been seen only in the feathertail glider. Our findings for the dLGN are of interest in the light of recent serological and taxonomic studies which suggest a close link between the feathertail glider and the honey possum.

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PubMed1989

Cell death in the inner and outer nuclear layers of the developing retina in the wallaby Setonix brachyurus (quokka).

We have examined the number and distribution of dying cells in the developing inner (INL) and outer (ONL) nuclear layers of sectioned quokka retinae (N = 31) from embryonic day (E)24 to postnatal day (P)192. Before birth, dying cells were seen in the optic fissure. Thereafter two major phases of cell death took place in the INL. The first phase was more pronounced within the vitread part with peak numbers of dying cells at P50. By contrast, during the second phase, cell death was more extensive in the sclerad portion; peak numbers of dying cells were recorded at P85 and P100 for the vitread and sclerad parts respectively. At these stages, photoreceptors were seen in the INL suggesting that these ectopic cells contribute to the pool of dying cells. The pattern of cell death broadly followed a central to peripheral sequence in the first phase but, in the second, was seen initially in mid-temporal retina and then became panretinal. Dying cells were seen in the ONL but in smaller numbers than in the INL. There was a peak of cell death at P26 which may represent death of mitotic cells at the ventricular surface. In the quokka, retinal cell genesis takes place in two phases (Harman and Beazley: Neuroscience 28:219-232, '89). The two major phases of cell death described here peak approximately 40 days after episodes of maximal cell genesis. These findings, together with data for the mouse, suggest that a biphasic pattern of cell genesis and cell death may be a feature of eutherian as well as marsupial retinal development.

Aging↗

PubMed1989

Expanded retinofugal projections to the dorsal lateral geniculate nucleus and superior colliculus after unilateral enucleation in the wallaby Setonix brachyurus, quokka.

We removed one eye of quokkas either neonatally, before retinal innervation of visual centres, or at 35-40 days postnatal, when projections overlap bilaterally and are more widespread than in the adult. Retinal projections to the dorsal lateral geniculate nucleus and superior colliculus at postnatal day 100 were demonstrated following anterograde transport of horseradish peroxidase. There were significant reductions in the size of the dorsal lateral geniculate nucleus and superior colliculus ipsilateral to the remaining eye. However, the extent of retinofugal projections was markedly expanded in comparison to the normal input from one eye. Unexpectedly, projections were expanded to similar extents in the two series of enucleated animals although ipsilateral labelling appeared more dense after neonatal enucleation. In controls, label was restricted to eye-specific regions but in enucleated animals there were no label-free zones. Nevertheless the alpha laminae remained distinct in the dorsal lateral geniculate nucleus of enucleated animals. Our findings suggest that binocular interactions are necessary for the segregation and refinement of visual projections but not for the formation of the alpha laminae.

Animals↗

PubMed1989

Retinal ganglion cell number is unchanged in the remaining eye following early unilateral eye removal in the wallaby Setonix brachyurus, quokka.

The expanded visual projections which develop after unilateral eye removal have been associated in some studies, but not in others, with the survival of more ganglion cells than normal in the remaining eye. We have addressed this issue using the small wallaby Setonix brachyurus, quokka. Moreover to determine whether more ganglion cells survive when the eye is removed at a very early stage, we have compared the effect of enucleations at two ages. These were within 3 days of birth, before optic fibres innervate visual centres, and at 35-40 days postnatal, when visual projections are exuberant. At 100 days postnatal, retinal ganglion cells were retrogradely labelled from primary visual centres and tracts with horseradish peroxidase, allowing 24 h for transport. Numbers of ganglion cells were similar between animals enucleated as neonates (X = 231,000, n = 3) and at 35-40 days postnatal (X = 218,000, n = 4). These results were comparable to those of controls (X = 227,000, n = 5). Distributions of ganglion cells were also essentially similar in experimental and control series. However, mean ganglion cell soma diameter was significantly greater than normal in both the area centralis and temporal retina after neonatal enucleation. Our results indicate that in enucleated quokkas increased ganglion cell numbers do not underlie enhanced retinofugal projections.

Animals↗

PubMed1989

Neurogenesis and cell death in the isthmic nuclei of the frog Limnodynastes dorsalis.

In Limnodynastes dorsalis, neurogenesis of the isthmic nuclei was determined by 3H-thymidine autoradiography. The nuclei developed in a rostroventral to caudodorsal direction. Peak neurogenesis occurred in midlarval life, resulting in the formation of a substantial portion of each nucleus. Cell death was evidenced by the presence of pyknotic cells within the nucleus in all except the earliest stages. Peak cell death occurred around metamorphic climax. It was during this period that a rim neuropil appeared that segregates within the nucleus the contralateral and ipsilateral components of the tecto-isthmo-tectal pathways underlying binocular vision. It is also a time just prior to the onset of electrophysiologically detectable ipsilateral visual responses. However, no systematic cell death was found. Therefore the role of cell death in the formation of the isthmic nuclei remains obscure, although it may be involved in the establishment of tecto-isthmo-tectal connections.

Animals↗

PubMed1989

Prevention of optic nerve regeneration in the frog Hyla moorei transiently delays the death of some ganglion cells.

We have previously reported that a proportion of ganglion cells die during optic nerve regeneration in the adult frog Hyla moorei (Humphrey and Beazley, '85). Here we assess the effect of preventing optic nerve regeneration on this cell loss. The optic nerve was crushed unilaterally and regeneration was allowed to progress unimpeded in one experimental series but was prevented by ligating or capping the nerve in another. We estimated total cell numbers in the ganglion cell layer from cresyl-stained wholemounts, comparing each experimental retina with its unoperated partner. At 70-78 days postcrush, mean cell numbers had fallen by 31.5% for frogs with unimpeded regeneration (N = 9), a significantly greater reduction than the 21.5% (N = 8) loss for the impeded regeneration series (p less than 0.001). Thereafter, cell numbers were stable for frogs with unimpeded regeneration. Cell death continued in the series with impeded regeneration, and losses exceeded those of frogs with unimpeded regeneration from 110 days postcrush. When regeneration was impeded, ganglion cell somas underwent an intense cell soma reaction and became arranged in rows radiating from the optic nerve head. Our findings indicate that some ganglion cells are transiently spared when regeneration of their axons is prevented. The abnormally extensive contacts formed between somas may delay ganglion cell loss. However, the eventual death of most ganglion cells shows them to be target-independent in the long term.

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PubMed1989

Generation of retinal cells in the wallaby, Setonix brachyurus (quokka).

We have examined the generation of retinal cells in the wallaby, Setonix brachyurus (quokka). Animals received a single injection of tritiated thymidine between postnatal days 1-85 and retinae were examined at postnatal day 100. Retinae were sectioned, processed for autoradiography and stained with Cresyl Violet. Ganglion cells were labelled by injection of horseradish peroxidase into the optic tracts and primary visual centres. Other cells were classified according to their morphology and location. Retinal cell generation takes place in two phases. During the first phase, which concludes by postnatal day 30, cells destined to lie in all three cellular layers of the retina are produced. In the second phase, which starts by postnatal day 50, cell generation is almost entirely restricted to the inner and outer nuclear layers. Cells produced in the first phase are orthotopic and displaced ganglion cells, displaced and orthotopic amacrine cells, horizontal cells and cones. Glia in the ganglion cell layer, orthotopic amacrine cells, bipolar and horizontal cells. Muller glia, and rods are generated in the second phase. Cells became heavily labelled with tritiated thymidine in the central retina before postnatal day 7, over the entire retina (panretinal) by postnatal day 7 and from postnatal day 18, only in the periphery. The second phase of cell generation is initiated at P50, in a region extending from the optic nerve head to mid-temporal retina. Subsequently, cells are generated in annuli, centred on mid-temporal retina, which are seen at progressively more peripheral locations. Therefore, cell addition to the inner and outer nuclear layers continues for longer in peripheral than in mid-temporal retina. We suggest that such later differential cell addition to the inner and outer nuclear layers contributes to an asymmetric increase in retinal area. This non-uniform growth presumably results in more expansion of the ganglion cell layer peripherally than in mid-temporal retina and may play a role in establishing density gradients of ganglion cells.

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