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Retina, Iris, and Ciliary Body

The outer layer of the optic cup, which is characterized by small pigment granules,
is known as the pigmented layer of the retina (Figs. 17.3, 17.4, and 17.7).
Development of the inner (neural) layer of the optic cup is more complicated.
The posterior four-fifths, the pars optica retinae, contains cells bordering the
intraretinal
space (Fig. 17.3) that differentiate into light-receptive elements, rods
and cones (Fig. 17.5).
Adjacent to this photoreceptive layer is the mantle layer,
which, as in the brain, gives rise to neurons and supporting cells, including the
outer nuclear layer, inner nuclear layer, and ganglion cell layer (Fig. 17.5).
On the surface is a fibrous layer that contains axons of nerve cells of the deeper
layers. Nerve fibers in this zone converge toward the optic stalk, which develops
into the optic nerve (Figs. 17.3 and 17.5). Hence, light impulses pass through
most layers of the retina before they reach the rods and cones.
The anterior fifth of the inner layer, the pars ceca retinae, remains one cell
layer thick. It later divides into the pars iridica retinae, which forms the inner
layer of the iris, and the pars ciliaris retinae, which participates in formation
of the ciliary body (Fig. 17.6 and 17.7).
Meanwhile, the region between the optic cup and the overlying surface
epithelium is filled with loose mesenchyme (Figs. 17.3, 17.4, and 17.7). The
sphincter and dilator pupillae muscles form in this tissue (Fig. 17.6). These
muscles develop from the underlying ectoderm of the optic cup. In the adult,
the iris is formed by the pigment-containing external layer, the unpigmented
internal layer of the optic cup, and a layer of richly vascularized connective
tissue that contains the pupillary muscles (Fig. 17.6).
The pars ciliaris retinae is easily recognized by its marked folding
(Figs. 17.6B and 17.7). Externally it is covered by a layer of mesenchyme that
forms the ciliary muscle; on the inside it is connected to the lens by a network
of elastic fibers, the suspensory ligament or zonula (Fig. 17.7). Contraction of
the ciliary muscle changes tension in the ligament and controls curvature of
the lens.

Retina
The retina develops from the walls of the optic cup, an outgrowth of the forebrain (see Figs.
18-1 and 18-2). The outer, thinner layer of the optic cup becomes the retinal pigment
epithelium (pigmented layer of retina), and the inner, thicker layer differentiates into the
neural retina (neural layer of retina).

During the embryonic and early fetal periods, the two retinal layers are separated by an
intraretinal space (see Fig. 18-4), which is the original cavity of the optic cup. Before birth,
this space gradually disappears as the two layers of the retina fuse (see Fig. 18-8D), but this
fusion is not firm; hence, when an adult eyeball is dissected, the neural layer is often
separated from the pigment layer. Because the optic cup is an outgrowth of the forebrain, the
layers of the optic cup are continuous with the wall of the brain (see Fig. 18-1H).
Under the influence of the developing lens, the inner layer of the optic cup proliferates to
form a thick neuroepithelium (see Fig. 18-4). Subsequently, the cells of this layer
differentiate into the neural retina, the light-sensitive region of the optic part of the retina.
This region contains photoreceptors (rods and cones) and the cell bodies of neurons (e.g.,
bipolar and ganglion cells).
Fibroblast growth factor signaling regulates retinal ganglion cell differentiation. Because the
optic vesicle invaginates as it forms the optic cup, the neural retina is "inverted," that is, light-
sensitive parts of the photoreceptor cells are adjacent to the retinal pigment epithelium. As a
result, light must pass through the thickest part of the retina before reaching the receptors;
however, because the retina overall is thin and transparent, it does not form a barrier to light.
The axons of ganglion cells in the superficial layer of the neural retina grow proximally in the
wall of the optic stalk to the brain (see Figs. 18-3 and 18-4). As a result, the cavity of the
optic stalk is gradually obliterated as the axons of the many ganglion cells form the optic
nerve (see Fig. 18-3F).
Myelination of optic nerve fibers is incomplete at birth. After the eyes have been exposed to
light for approximately 10 weeks, myelination is complete, but the process normally stops
short of the optic disc, where the optic nerve enters the eyeball. Normal newborn infants can
see, but not too well; they respond to changes in illumination and are able to fixate points of
contrast. Visual acuity has been estimated to be in the range of 20/400. At 2 weeks of age, the
infants show a more sustained interest in large objects.

Lens
The lens develops from the lens vesicle, a derivative of the surface ectoderm (see Fig. 18-1).
The anterior wall of this vesicle, composed of cuboidal epithelium, becomes the subcapsular
lens epithelium (see Fig. 18-8C). The nuclei of the tall columnar cells forming the posterior
wall of the lens vesicle undergo dissolution. These cells lengthen considerably to form highly
transparent epithelial cells, the primary lens fibers. As these fibers grow, they gradually
obliterate the cavity of the lens vesicle (see Figs. 18-8A to C and 18-10).
The rim of the lens is known as the equatorial zone because it is located midway between
the anterior and posterior poles of the lens (Fig. 18-11). The cells in the equatorial zone are
cuboidal; as they elongate, they lose their nuclei and become secondary lens fibers. These
new lens fibers are added to the external sides of the primary lens fibers. Although secondary
lens fibers continue to form during adulthood and the lens increases in diameter, the primary
lens fibers must last a lifetime.
Choroid, Sclera, and Cornea

At the end of the fifth week, the eye primordium is completely surrounded by
loose mesenchyme (Fig. 17.3). This tissue soon differentiates into an inner layer

comparable with the pia mater of the brain and an outer layer comparable with
the dura mater. The inner layer later forms a highly vascularized pigmented
layer known as the choroid; the outer layer develops into the sclera and is
continuous with the dura mater around the optic nerve (Fig. 17.7).
Differentiation of mesenchymal layers overlying the anterior aspect of the
eye is different. The anterior chamber forms through vacuolization and splits
the mesenchyme into an inner layer in front of the lens and iris, the iridopupillary
membrane, and an outer layer continuous with the sclera, the substantia
propria of the cornea (Fig. 17.7). The anterior chamber itself is lined by flattened
mesenchymal cells. Hence, the cornea is formed by (a) an epithelial layer
derived fromthe surface ectoderm, (b) the substantia propria or stroma, which
is continuous with the sclera, and (c) an epithelial layer, which borders the anterior
chamber. The iridopupillary membrane in front of the lens disappears
completely, providing communication between the anterior and posterior eye
chambers.
Vitreous Body
Mesenchyme not only surrounds the eye primordium from the outside but also
invades the inside of the optic cup by way of the choroid fissure. Here it forms
the hyaloid vessels, which during intrauterine life supply the lens and form the
vascular layer on the inner surface of the retina (Fig. 17.7). In addition, it forms
a delicate network of fibers between the lens and retina. The interstitial spaces
of this network later fill with a transparent gelatinous substance, forming the
vitreous body (Fig. 17.7). The hyaloid vessels in this region are obliterated and
disappear during fetal life, leaving behind the hyaloid canal.
Optic Nerve
The optic cup is connected to the brain by the optic stalk, which has a groove,
the choroid fissure, on its ventral surface (Figs. 17.2 and 17.3). In this groove
are the hyaloid vessels. The nerve fibers of the retina returning to the brain
lie among cells of the inner wall of the stalk (Fig. 17.8). During the seventh
week, the choroid fissure closes, and a narrow tunnel forms inside the optic
stalk (Fig. 17.8B). As a result of the continuously increasing number of nerve
fibers, the inner wall of the stalk grows, and the inside and outside walls of the
stalk fuse (Fig. 17.8C ). Cells of the inner layer provide a network of neuroglia
that support the optic nerve fibers.
The optic stalk is thus transformed into the optic nerve. Its center contains
a portion of the hyaloid artery, later called the central artery of the retina. On
the outside, a continuation of the choroid and sclera, the pia arachnoid and
dura layer of the nerve, respectively, surround the optic nerve.