The earliest embryonic stage at which ocular structures can be differentiated from the rest of the fetus is the embryonic plate stage. The site of the eye is indicated by flattened areas on the both sides of the anterior of the neural groove. The edges of the neural groove (2.5 mm stage or 2 week stage) thicken to form the neural folds. The folds fuse to form the neural tube, which sinks into the underlying mesoderm and detaches from surface ectoderm. Before the anterior end of the neural tube is completely closed, buds of neural ectoderm on either side to form the spherical optic vesicle (4 mm or 3 week stage). The vesicles are connected to the forebrain by the optic stalks. At the 4 mm stage, a thickening of the surface ectoderm, the lens plate, begins to form just opposite the ends of the optic vesicles. OPTIC CUP STAGE The optic vesicle invaginates to produce the optic cup, so that the original outer wall of the optic vesicle becomes approximated to its inner wall. Invagination of the lower surface of the optic stalk and the optic vesicle occurs simultaneously, creating a groove known as the choroidal (fetal) fissure. At the same time, the lens plate also invaginates to form first a cup and then a hallow sphere known as the lens vesicle. By the 9 mm (4 week) stage, the lens vesicle separates completely from the surface ectoderm to lie free in the rim of the optic cup. The choroidal fissure permits entrance into the optic stalk of the vascular mesoderm which eventually forms the hyaloids system. As invagination becomes complete, the choroidal fissure narrows until it becomes completely closed (13 mm or 6 week stage), leaving one small permanent opening at the anterior end of optic stalk through which pass the hyaloids artery until the 100 mm (4 month) stage and the central retinal artery and vein thereafter. At this point, the ultimate general structure of the eye has been determined. Further the devolepment consist of the differentiation into individual structures. In general, differentiation occurs relatively more rapidly in the posterior than in the anterior segment early in gestation and more rapidly in the anterior segment later in gestation.
EMBRYONIC ORIGINS OF INDIVIDUAL EYE STRUCTURES Surface Ectoderm Lens; epithelium of cornea, conjunctiva, lacrimal gland and drainage system; vitreous (mesoderm also contributes to vitreous). Neural Ectoderm Vitreous, retina; epithelium of iris, ciliary body, and retina; papillary muscle sphincter and dilator, optic nerve. Mesoderm Sclera; stroma of cornea, conjunctiva, iris, ciliary body, choroid; extraocular muscles, lids (except epithelium and conjunctiva), hyaloid system (gone by birth), sheaths of the optic nerve; connective tissue and blood supply of eye, bony orbit, and vitreous.
EMBRYOLOGY OF SPECIFIC STRUCTURES Lens Soon after the lens vesicle lies free in the rim of the optic cup (13 mm or 6 week stage), the cells of its posterior wall elongate, encroach on the empty cavity, and finally fill it in (26 mm or 7 week stage). At about this stage (13 mm or 7 week stage), a hyaline capsule is secreted by the lens cells. Secondary lens fibers elongate from the equatorial region and grow forward under the subcapsular epithelium, which remains as a single layer of cuboid epithelial celss, and backward under the lens capsule. These fibers meet to form the lens sutures (upright Y anteriorly and inverted Y posteriorly), which are complete by the seventh month. (this growt and proliferation of the secondary lens fibers continues at a decreasing rate throughout life; the lens therefore continues to enlarge slowly, causing compression of the lens fibers).
Retina The outer layer of the optic cup remains as a single layer and becomes the pigment epithelium of the retina. Pigmentation begins at the 10 mm (5 week) stage. The inner layer undergous a complicated differentiation into the other 9 layers of the retina. This occurs slowly throughout gestation. By the seventh month, the outermost cell layer (consisting of the nuclei of the rods and cones) is present as well as the bipolar, amacrine, and ganglion cells and nerve fibers. The macular region is thicker than the rest of the retina until the eight month, when macular depression begins to develop. Macular devolepment is not complete until 6 months after birth. Optic Nerve The axons of the ganglion cell layer of the retina form the inner nerve fiber layer. The fibers slowly form the optic stalk and then the optic nerve (26 mm or 7 week stage). Mesodermal elements enter the surrounding tissue to form the vascular septa of the nerve. Medullation extends from the brain peripherally down the optic nerve, and the birth has reached the lamina cribrosa. Medullation is completed by age 3 months.
Iris & Ciliary body During the third month (50 mm stage), the rim of the optic cup grows forward in front of the lens as a double row of epithlelium and lies posterior to mesoderm, which becomes the stroma of the iris. These 2 epithelial layers become pigmented in the iris, whereas only outer layer is pigmented in the ciliary body. Folds appear in the epithelial layers of the ciliary body; mesoderm grows into this fold to form the ciliary processes. By the fifth month (150 mm stage), the sphincter muscle of the pupil is developing from a bud of nonpigmented epithelium derived from the anterior epithelial layer of the iris near the papillary margin. Soon after the sixth month, the dilator muscle appears in the anterior epithelial layer near the ciliary body.
Choroid At the 6 mm (3 1 / 2 week) stage, a network of capillaries encircles the optic cup and develops into the choroid. By the 13 mm (6 week) stage, the outer neural epithelial layer has secreted Bruchs membrane. By the third month, the intermediate and large venous channels of the choroid are developed and drain in the vortex veins to exit from the eye.
Vitreous A. First stage: (primary vitreous, 4.5-13 mm or 3-6 week stage). At about the 4.5 mm stage, fibrils grow in from the inner layer of the optic vesicle to join elements from the lens vesicle which along with some mesoderm fibrils as sociated the hyaloid artery form the primary vitreous. This stage ends as the lens capsule appears, precluding any further lens participation in vitreous formation. The primary vitreous does not atrophy and ultimately lies just behind the posterior pole of the lens as the hyaloids canal. B. Second Stage: (Secondary vitreous, 13-65 mm or 3-10 week stage). Mllers fibers of the retina become continuous with vitreous fibrils, so that the secondary vitreous is mainly derived from retinal ectoderm. The hyaloid system develops a set of vitreous vessels as well as vessels on the lens capsule surface (tunica vasculosa lentis). The hyaloid system is the its height at 40 mm and then atrophies from posterior to anterior. C. Third Stage: tertiary vitreous, 65 mm or 10 weeks on). During the third month, the marginal bundle of Drualt is forming. This consists of vitreous fibrillar condensations extending from the future ciliary epithelium of the optic cup to the equator of the lens, which is well developed by the 100 mm or 4 month stage. The hyaloid system atrophies completely during this stage.
Blood Vessels Long ciliary arteries bud off from the hyaloid at the 16 mm (6 week) stage and anastomose around the optic cup margin with the major circle of the iris by the 30 mm (7 week) stage. The hyaloid system (see vitreous) atrophies completely by the eight month. The hyaloid artery gives rise to the central retina artery and its branches (100 mm or 4 month stage). Buds begin to grow in to the retina and develop the retinal circulation, which reaches the ora serrata at 8 months. The branches of the central retinal vein develop simultaneously.
Cornea The epithelium is derived from surface ectoderm, whereas the rest of the cornea comes mesodermal structures. The earliest differentiation is seen in at about the 12 mm (5 week) stage, when endothelial cells appear.