We have studied the rise and fall in the number of axons in the optic nerve of fetal and neonatal cats in relation to changes in the ultrastructure of fibers, and in particular, to the characteristics and spatiotemporal distribution of growth cones and necrotic axons. Axons of retinal ganglion cells start to grow through the optic nerve on the 19th day of embryonic development (E-19). As early as E-23 there are 8,000 fibers in the nerve close to the eye. Fibers are added to the nerve at a rate of approximately 50,000 per day from E-28 until E-39--the age at which the peak population of 600,000-700,000 axons is reached. Thereafter, the number decreases rapidly: About 400,000 axons are lost between E-39 and E-53. In contrast, from E-56 until the second week after birth the number of axons decreases at a slow rate. Even as late as postnatal day 12 (P-12) the nerve contains an excess of up to 100,000 fibers. The final number of fibers--140,000-165,000--is reached by the sixth week after birth. Growth cones of retinal ganglion cells are present in the optic nerve from E-19 until E-39. At E-19 and E-23 they have comparatively simple shapes but in older fetuses they are larger and their shapes are more elaborate. As early as E-28 many growth cones have lamellipodia that extend outward from the core region as far as 10 microns. These sheetlike processes are insinuated between bundles of axons and commonly contact 10 to 20 neighboring fibers in single transverse sections. At E-28 growth cones make up 2.0% of the fiber population; at E-33 they make up about 1.0%; from E-36 to E-39 they make up only 0.3% of the population. Virtually none are present in the midorbital part of the nerve on or after E-44. At all ages growth cones are more common at the periphery of the nerve than at its center. This central-to-peripheral gradient increases with age: at E-28 the density of growth cones is two times greater at the edge than at the center but by E-39 the density is four to five times greater. Necrotic fibers are observed as early as E-28 in all parts of the nerve. Their axoplasm is dark and mottled and often contains dense vesiculated structures.(ABSTRACT TRUNCATED AT 400 WORDS)
In order to investigate the development of luminance and chromatic temporal contrast sensitivity functions (tCSFs), we obtained chromatic and luminance contrast thresholds from individual 3- and 4-month old infants, and compared them to previously obtained functions in adults. Stimuli were moving sinusoidal gratings of 0.27 cyc/deg, presented at one of five temporal frequencies: 1.0, 2.1, 4.2, 9.4 or 19 Hz (corresponding speeds: 3.8, 7.7, 15, 34, 69 deg/s). Previous studies, including our own, have shown that adult tCSFs are bandpass for luminance stimuli (peaking at 5-10 Hz), yet lowpass for chromatic stimuli (sensitivity falling at > 2 Hz), and that the two functions cross one another near 4-5 Hz when plotted in terms of cone contrast. In the present study, we find that the shapes and peaks of the luminance tCSF in both 3- and 4-months-olds appear quite similar to those of adults. By contrast, chromatic tCSFs in infants are markedly different from those of adults. In agreement with our earlier report (Dobkins, K. R., Lia, B., & Teller, D. Y. (1997). Vision Research, 37(19), 2699-2716), the chromatic function in 3-month-olds is rather flat, lacking the sharp high temporal frequency fall-off characteristic of the adult function. In addition, the luminance tCSF in 3-month-olds is elevated above the chromatic tCSF, and the two functions do not exhibit an adult-like cross-over within the range of temporal frequencies tested. By 4 months of age, substantial development of chromatic contrast sensitivity takes place at the lowest temporal frequencies. Although still immature, the 4-month-old chromatic tCSF has begun to adopt a more adult-like shape. In addition, similar to adults, luminance and chromatic tCSFs in 4-month-olds cross one another near 5 Hz. In adults, magnocellular (M) and parvocellular (P) pathways are thought to underlie the bandpass luminance and lowpass chromatic tCSF, respectively (e.g. Lee, B. B., Pokorny, J., Smith, V. C., Martin, P. R., & Valberg, A. (1990). Journal of the Optical Society of America (a), 7(12), 2223-2236). Based on this correspondence between psychophysical and neural responses in adults, our results suggest that the relatively slow development of the chromatic tCSF in infants may reflect immature chromatic responses in the P pathway and/or reliance on chromatic responses originating in the M pathway.
A fundamental feature of the mammalian visual system is the nonuniform distribution of ganglion cells across the retinal surface. To understand the ontogenetic processes leading to the formation of retinal ganglion cell topography, changes in the regional density of these neurons were studied in relation to ganglion cell loss and the pattern of retinal growth in the fetal cat. Midway through the gestation period, the density of these neurons was only two to three times greater in the area centralis than in the peripheral retina, whereas shortly before birth this central-to-peripheral difference was nearly 20-fold. Age-related changes in the ganglion cell distribution were found not to correspond in time or magnitude to the massive loss of ganglion cells that occurs during prenatal development. Rather, the formation of ganglion cell density gradients can be accounted for by unequal expansion of the growing fetal retina-peripheral regions expand more than the central region, thereby diluting the peripheral density of ganglion cells to a greater degree. Nonuniform growth, in conjunction with differential periods of neurogenesis of the different types of retinal cells, appears to be a dominant factor regulating overall retinal topography. These results suggest that the differential regional expansion of the fetal retina underlies the formation of magnification factors in the developing visual system.
The development of the partial decussation pattern in the primate retina was studied in fetal rhesus monkeys of known gestational ages. Retinal ganglion cells with either crossed or uncrossed projections were identified by labeling with HRP following unilateral injections of this tracer into the optic tract. At all fetal ages, very few cells (less than 0.5% of the total ganglion cell population) were found to project to the inappropriate hemisphere. The nasotemporal overlap zone, defined as the retinal region along the vertical meridian containing cells with either crossed or uncrossed projections, also appeared equivalent to that described for the adult animal. A temporal offset in the decussation pattern of large ganglion cells, similar to that of the mature retina, could be recognized as early as 50 d before birth. These results indicate that an adultlike retinal decussation pattern is evident in the fetal primate at a stage when projections from the 2 eyes are completely intermingled within retinorecipient nuclei, and prior to the onset of retinal ganglion cell loss. Moreover, the primate visual system exhibits a degree of precision in the specification of the nasotemporal division unrivaled among the mammalian species studied to date. The developmental specificity evident in the decussation pattern of the fetal rhesus monkey appears to reflect the specialized organization of this primate's retina for binocular focal vision.
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