Embryonic Development of the Eye

The vertebrate eye is derived from both ectodermal and mesodermal primordial sources in the cephalic region of the embryo. Mechanical influences are of the utmost importance in embryonic eye development to ensure the proper alignment of tissue in the visual pathway.

Initiation
Modern research indicates that the action of a master control gene initiates eye development. This gene, called eyeless in Drosphila, small eye (Pax-6) in the mouse, and Andridia in humans, acts as a transcription factor that sets off a cascade of other genes that are involved in later stages of eye development. Neural ectoderm is induced by underlying chordamesoderm.

Primary Optic Vesicle and Optic Cup

The first morpological sign of eye formation is the evagination of the wall of the diencephalon which appear as shallow concavities that deepen into the primary optic vesicles. As development continues, the distal portion of this single-walled vesicle flattens and then invaginates toward the vesicle's ventral margin to form the doublewalled optic cup. The thinner, outer layer of the wall is eventually converted into the pigment layer of the retina. The inner layer thickens and ultimately becomes the sensory layer of the retina. When the vesicle invaginates to form the optic cup, a gap in the wall's continuity is formed, called the choroid fissure. This fissure will partially envelope the hyloid artery, a branch of the internal carotid arterly, which supplies blood to many embryonic structires of the eye. The fissure also extends along part of the optic stalk which later develops into the optic nerve when synaptic connections are established with the brain.

Development of the Lens

72 Hour Chick Embryo

A= Lens Vesicle B= Retina, black line to the left is pigment layer C=Optic Stalk

As invagination is occuring to form the optic cup, the lens placode is formed by the thickening of the overlying head ectoderm. The inductive influence ot the opetic vesicle causes the future lens ectoderm to thicken and invaginate, forming a lens vesicle. This lens vesicle undergoes an asymmetrical development in which the deeper cells elongate into lens fibers. The cells of the outer pole keep their low epithelial configuration. Eventually, the lens fibers elongate enough to moke contact with the lens epithelium. The original cavity of the lens vesicle is reduced to a tiny slit. Cytodifferentiation within the lens equates to the transformation of dividing epthelial cells from the lens's outer pole into elongated postmitotic cells in the lens's main body. Lens fibers elongate fron epithelial cells in an equatorial zone that surrounds the entire lens. These changes prepare for the formation of large amounts of crystallins within the lens fibers. Crystallins are structural proteins translated from a long-lived mRNA in mitotically inactive cells. Today, scientists are examining evidence of a mechanism that ensures continuous embryological lens allignment.

Iris and Ciliary Apparatus

A=Pigment Layer of the Retina; B=Sensory Layer of the Retina; C=Optic Stalk; D=Lens Vesicle

The lens settles back into the optic cup as its size increases. The overlapping part of the optic cup forms the iris's epithelial portion. The iris's muscular portion is unusaul in that it arises not from mesoderm but from neurectoderm. The ciliary body's

muscular portion arises from loosely aggregated mesenchyme outside the epithelial layer. The amount of intraocular pressure must be correct in order for the ciliary body to develop normally.

Choroid Coat and Sclera

Mesenchymal cells outised the optic coat amass early in a concentration zone and react to an inductive influence from the optic cup, forming the inner, differentiated, vascular choroid coat. Oustide the choroid coat, other mesenchymal cells similarly infuenced form the sclera,composed of dense fibrous connective tissue.

Cornea
Ordinary surface ectoderm is stimulated to transform into corneal epithelium by an inductive influence from the lens vesicle and optic cup. This first corneal epithelium is two cells thick, and the Golgi apparatus of these cells shift toward the basal surface, froming the primary stroma. Corneal endothelial cells, which arise from mesodermal mesemchyme associated with blood vessels surrounding the lip of the optic cup, migrate over the inner surface of the primary stroma. These cells continue to form a complete inner corneal lining, and synthesize a large aMount of hyaluronic acid which is secreted into the primary stroma, causing the stroma to swell. This swelling signals a new phase in which fibroblasts settle. Large amounts of hyaluronidase is next seen, presumably synthesized by the fibroblasts. The stroma, now populated by fibroblasts, is referred to as the secondary stroma. The corneal stroma now dehydrates, due to thyroxine sent by the develping thyroid gland, in order to make the cornea transparent.

Retina
The outer wall of the optic cup forms the pigment layer of the retina. Cells of the inner wall of the optic cup, however, differentiate into neural elements, forming the neural (sensory) layer of the retina. Radial polarity is established first by the fixation of the nastotemporal axis and then the dorsoventral axis. The mature neural retina is compsed of three layers of ganglion, bipolar cells, and rods and cones, each of which is respectively further from the center of the eyeball. Embryonically, the ganglion layer, being the closest to the center of the eyeball, develops first, and the remaining layers differentiate outward. This differentiation is accompanied by waves of cell death. The reason for this is not yet fully understood. The neurons of the ganglion layer send out axons which grow out into the ventral wall of the optic stalk toward the brain's optic centers.

The retina of the eye comes from the neural tube.

The optic vesicle is derived from the anterior part of the neural tube, a region called the diencephalon. Genes like pax6 (Figure 1, arrow) specify this region of the developing nervous system to become the retina. After the evagination of the optic vesicles from the neural tube, the optic vesicle develops into an optic cup, and ultimately into a neural retina, the pigmented epithelium (RPE), the ciliary epithelium, and the iris (Figure 2). There is also a small zone of cells that lies between the retina and the ciliary epithelium in non-mammalian vertebrates, called the ciliary marginal zone, or CMZ (red), that contains retinal stem cells, which produce new neurons and glia throughout life in fish and some amphibians. Although derived from the same region of the neural tube, these tissues are quite distinct; for example, the neural retina is a multilayered structure containing the photoreceptors and neurons necessary for vision, while the RPE is a single layer of non-neuronal, pigment cells, necessary for support of the retinal photoreceptors. The anterior part of the optic vesicle normally forms the neural retina, while the posterior part develops as the RPE. These different cell fates are not immediately fixed, however, and there is a period of development during which the fates of these two tissues can be reversed. For chicks and mammals, the fate of these cells becomes restricted to either the RPE or neural retinal cell fateshortly after the formation of the optic cup; however, in amphibians this cell fate decision is not fixed during development and the RPE can regenerate neural retina after damage (see RetinalRegeneration in Amphibians).

Eyelids and Conjunctiva

Eyelids develop as folds of skin growing back over the cornea, which ultimately fuse. Later in fetal development, the eyelids once again separate. The conjunctival sac, the space between the front of the eyball and the eyelids, develop from multipal epithelial buds. The conjunctival sac contains the lacrimal glands, or tear ducts.

Eye Position

As facial structure develops, the eyes that began to form on either side of a human embryo's head are carried forward. This allows for the convergence of the optical axes.

In vitro system to study optic vesicle patterning.

To investigate the factors that pattern the optic vesicle into these various domains, we developed an explant culture preparation of the optic vesicle. Optic vesicles from E1.5 chick embryos (H&H stages 9-10) were removed and placed into tissue culture (Figure 3). The vesicles retained the associated ectoderm and surrounding mesenchymal cells. Under control conditions, the vesicles developed into optic cups after 24 hrs. The three layers of the cup, the pigment epithelium neural retina and lens can all be identified in the sections through the explants. In addition, immunohistochemistry and in situ hybridization for neural and RPE specific antigens confirm appropriate development of the neural and RPE layers in the explants.

FGF causes retinal duplication

When optic vesicles are maintained in medium containing FGF1 or FGF2, the RPE layer never develops and instead, both layers of the developing optic cup become neural retina. Antibodies against neural antigens, such as neurofilament (Figure 4) and neuron-specific tubulin, now label both layers of the vesicle (nr, and pe are labeled; Figure 4). So exogenous FGF2 is sufficient to cause the entire optic vesicle to form neural retinal tissue, and block the formation of the RPE. We have also found that experimentally inhibiting the FGF signaling pathway prevents neural retina formation, but does not affect the development of the RPE (Pittack et al, 1997). So we conclude that FGF signaling is both necessary and sufficient for the development of the neural retinal domain of the optic vesicle and optic cup.

Extraocular mesenchyme produces a signal that promotes the differentiation of the pigmented epithelium (RPE).
To determine what molecular signals pattern the other major domain of the optic vesicle-the RPE-Sabine Fuhrmann took a similar approach when she was a postdoctoral fellow in the lab. Classical studies in frog embryos indicated that the ectomesenchyme (the mesenchymal tissue surrounding the developing optic vesicle) was critical for the development of the RPE. Sabine Fuhrmann tested this idea for the chick embryo in the following way. First she dissected the optic vesicle along with the surrounding tissues, but then she removed the mesenchymal cells. When the optic vesicles without the extraocular mesenchyme were cultured, their RPE development was inhibited. What are the molecules released by the extraocular mesenchyme that are responsible for inducing RPE from the optic vesicle? She tested several different types of known developmental signaling molecules, including many different TGF-beta related molecules. She found that of all the factors that tested, activin was the most effective at replacing the extraocular mesenchyme in promoting the differentiation of the RPE. These experiments have led us to the following working model of how the neural retina and the RPE domains of the optic vesicle are patterned during eye development. An FGF signal, either from the cells of the lens ectoderm, or from the optic vesicle cells themselves, promotes the distal part of the optic vesicle to develop as neural retina. An

opposing signal, activin, from the mesenchyme surrounding the optic vesicle promotes the development of the RPE. Full details of this model and additional experimental details can be found in:

Transverse section of head of chick embryo of forty-eight hours incubation.

Transverse section of head of chick embryo of fifty-two hours incubation, showing the lens and the optic cup. The eye begins to develop as a pair of optic vesicles on each side of the forebrain at the end of the 4th week of pregnancy. Optic vesicles are outgrowings of the brain which make contact with the surface ectoderm and this contact induces changes necessary for further development of the eye. Through a groove at the bottom of the optic vesicle known as choroid fissure the blood vessels enter the eye. Several layers such as the neural tube, neural crest, surface ectoderm, and mesoderm contribute to the development of the eye.[1]

Contents

1 Sequential inductions o 1.1 Neural tube ectoderm (Neuroectoderm) o 1.2 Surface Ectoderm o 1.3 Neural crest o 1.4 Mesoderm 2 Developmental cascade 3 Responsivity of head epidermis 4 Regulation and inhibition 5 Additional images 6 References 7 External links

Sequential inductions[edit]

This development is an example of sequential inductions where the organ is formed from three different tissues: Neural tube ectoderm (Neuroectoderm)[edit] First, there is an outpocketing of the neural tube called optic vesicles. Development of the optic vesicles starts in the 3-week embryo, from a progressively deepening groove in the neural plate called the optic sulcus. As this expands, the rostral neuropore (the exit of the brain cavity out of the embryo) closes and the optic sulcus and the neural plate becomes the optic vesicle.[2] Neuroectoderm gives rise to the following compartments of the eye:

Retina Epithelial lining of Ciliary body and Iris Optic nerves vitreous

Neuroectoderm
From Wikipedia, the free encyclopedia Neuroectoderm

Latin Precursor

epithelium tubi neuralis, neuroectoderma, epithelium tubae neuralis ectoderm

Gives rise to Code

neural tube, neural crest TE E5.15.1.0.0.0.1

Neuroectoderm (or neural ectoderm or neural tube epithelium) is the term for ectoderm which receives Bone Morphogenetic Protein-inhibiting signals from proteins such as noggin, which leads to the development of the nervous system from this tissue. After recruitment from the ectoderm, the neuroectoderm undergoes three stages of development: transformation into the neural plate, transformation into the neural groove (with associated neural folds), and transformation into the neural tube. After formation of the tube, the brain forms into three sections; the hindbrain, the midbrain, and the forebrain. The types of neuroectoderm include:

Neural crest

pigment cells in the skin ganglia of the autonomic nervous system dorsal root ganglia. facial cartilage spiral septum of developing heart ciliary body of the eye adrenal medulla

Neural tube

brain (rhombencephalon, mesencephalon and prosencephalon) spinal cord and motor neurons retina posterior pituitary

Retina

From Wikipedia, the free encyclopedia

Not to be confused with retinal.

Retina

Right human eye cross-sectional view. CourtesyNIH National Eye Institute. Many animals have eyes different from the human eye.

Gray's

subject #225 1014

Artery

central retinal artery

MeSH

Retina

Dorlands/Elsevier

Retina

Retina can also refer to the Kodak Retina camera and the Apple Retina Display. The vertebrate retina (/rtn/ RET-nuh, pl. retinae, /rtini/; from Latin rte, meaning "net") is a lightsensitive layer of tissue, lining the inner surface of the eye. The optics of the eye create an image of the visual world on the retina (through the cornea and lens) , which serves much the same function as the film in a camera. Light striking the retina initiates a cascade of chemical and electrical events that ultimately trigger nerve impulses. These are sent to various visual centres of the brain through the fibres of the optic nerve.

In vertebrate embryonic development, the retina and the optic nerve originate as outgrowths of the developing brain, so the retina is considered part of the central nervous system (CNS) and is actually brain tissue.[1][2] It is the only part of the CNS that can be visualized non-invasively.

The retina is a layered structure with several layers of neurons interconnected by synapses. The only neurons that are directly sensitive to light are the photoreceptor cells. These are mainly of two types: the rods and cones. Rods function mainly in dim light and provide black-and-white vision, while cones

support daytime vision and the perception of colour. A third, much rarer type of photoreceptor, the photosensitive ganglion cell, is important for reflexive responses to bright daylight.

Neural signals from the rods and cones undergo processing by other neurons of the retina. The output takes the form of action potentials in retinal ganglion cells whose axons form the optic nerve. Several important features of visual perception can be traced to the retinal encoding and processing of light.

Ciliary body
From Wikipedia, the free encyclopedia

Ciliary body

Schematic diagram of the human eye

Latin

corpus ciliare

Gray's

subject #225 1010

Artery

long posterior ciliary arteries

MeSH

Ciliary+Body

The ciliary body is the circumferential tissue inside the eye composed of the ciliary muscle and ciliary processes.[1] It is triangular in horizontal section and is coated by a double layer, the ciliary epithelium. This epithelium produces the aqueous humor.[2] The inner layer is transparent and covers the vitreous body, and is continuous from the neural tissue of the retina. The outer layer is highly pigmented,

continuous with the retinal pigment epithelium, and constitutes the cells of the dilator muscle. This double membrane is often regarded to be continuous with the retina and a rudiment of the embryological correspondent to the retina. The inner layer is unpigmented until it reaches the iris, where it takes on pigment. The retina ends at the ora serrata. The ciliary body is part of the uveal tract the layer of tissue which provides most of the nutrients in the eye. It extends from the ora serrata to the root of the iris.

There are three sets of ciliary muscles in the eye, the longitudinal, radial, and circular muscles. They are near the front of the eye, above and below the lens. They are attached to the lens by connective tissue called the zonule of Zinn, and are responsible for shaping the lens to focus light on the retina. The ciliary body receives parasympathetic innervation from the oculomotor nerve.

Functions[edit]

3D rendering of the eye. See related animation of ciliary body.

The ciliary body has three functions: accommodation, aqueous humor production and the production and maintenance of the lens zonules. It also anchors the lens in place. Accommodation essentially means that when the ciliary muscle contracts, the lens becomes more convex, generally improving the focus for closer objects. When it relaxes it flattens the lens, generally improving the focus for farther objects. One of the essential roles of the ciliary body is also the production of the aqueous humor, which is responsible for providing most of the nutrients for the lens and the cornea and involved in waste management of these areas.

Clinical significance[edit]
The ciliary body is the main target of drugs against glaucoma (apraclonidine) as it is responsible for aqueous humor production. Its inhibition leads to the lowering of aqueous humor production and causes a subsequent drop in the intraocular pressure.

Iris (anatomy)
From Wikipedia, the free encyclopedia

This article is about the Iris (anatomy). For other uses, see Iris (disambiguation).

Iris

The iris is the blue area. The other structures visible are the pupil center and the white sclerasurrounding the iris. The overlying cornea is pictured, but not visible, as it is transparent. Also pictured are the red-colored blood vessels within the sclera. These are easily visible on any person's eyes.

Schematic diagram of the human eye. (Iris labeled at upper left.)

Gray's

subject #225 1012

Artery

long posterior ciliary arteries

Nerve

long ciliary nerves, short ciliary nerves

MeSH

Iris

Dorlands/Elsevier

Iris

Medical portal Optometry portal

The iris (plural: irides or irises) is a thin, circular structure in the eye, responsible for controlling the diameter and size of the pupil and thus the amount of light reaching the retina. The color of the iris is often referred to as "eye color."
Contents
[hide]

1 Etymology 2 Embryology 3 General structure

o o o

3.1 Histological features 3.2 Anterior surface features 3.3 Posterior surface features

4 Eye color

o o

4.1 Genetic and physical factors determining iris color 4.2 Different colors in the two eyes

5 Conjunctivitis or "pink eye" 6 Alternative medicine 7 Graphics 8 See also 9 References 10 External links

Etymology[edit]
The word iris is derived from the Greek goddess of the rainbow, due to the many colours of the iris.

Embryology[edit]

The iris develops from the anterior two layers of an embryonic neuroectoderm structure called the optic cup. The optic cup also produces the iris sphincter and dilator muscles.

General structure[edit]
The iris consists of two layers: the front pigmented fibrovascular tissue known as a stroma and, beneath the stroma, pigmented epithelial cells. The stroma connects to a sphincter muscle (sphincter pupillae), which contracts the pupil in a circular motion, and a set of dilator muscles (dilator pupillae) which pull the iris radially to enlarge the pupil, pulling it in folds. The back surface is covered by a heavily pigmented epithelial layer that is two cells thick (the iris pigment epithelium), but the front surface has no epithelium. This anterior surface projects as the dilator muscles. The high pigment content blocks light from passing through the iris to the retina, restricting it to the pupil. [1] The outer edge of the iris, known as the root, is attached to the sclera and the anterior ciliary body. The iris and ciliary body together are known as the anterior uvea. Just in front of the root of the iris is the region referred to as the trabecular meshwork, through which the aqueous humour constantly drains out of the eye, with the result that diseases of the iris often have important effects on intraocular pressure and body provide a lesser secondary pathway for the aqueous humour to drain from the eye. The iris is divided into two major regions: 1. The pupillary zone is the inner region whose edge forms the boundary of the pupil. 2. The ciliary zone is the rest of the iris that extends to its origin at the ciliary body. The collarette is the thickest region of the iris, separating the pupillary portion from the ciliary portion. The collarette is a rudiment of the coating of the embryonic pupil.[1] It is typically defined as the region where the sphincter muscle and dilator muscle overlap. Radial ridges extend from the periphery to the pupillary zone, to supply the iris with blood vessels. The root of the iris is the thinnest and most peripheral.[2] The muscle cells of the iris are smooth muscle in mammals and amphibians, but are striated muscle in birds and reptiles. Many fish have neither, and, as a result, their irides are unable to dilate and contract, so that the pupil always remains of a fixed size.[3]

Histological features[edit]
From anterior (front) to posterior (back), the layers of the iris are:

Anterior limiting layer Stroma of iris Iris sphincter muscle Iris dilator muscle

Anterior pigment myoepithelium Posterior pigment epithelium

Anterior surface features[edit]

The Crypts of Fuchs are a series of openings located on either side of the collarette that allow the stroma and deeper iris tissues to be bathed in aqueous humor. Collagen trabeculae that surround the border of the crypts can be seen in blue irides.

The midway between the collarette and the origin of the iris. These folds result from changes in the surface of the iris as it dilates.[citation needed]

Crypts at the base of the iris are additional openings that can be observed close to the outermost part of the ciliary portion of the iris.[2]

Posterior surface features[edit]

The Radial contraction folds of Schwalbe are a series of very fine radial folds in the pupillary portion of the iris extending from the pupillary margin to the collarette. They are associated with the scalloped appearance of the pupillary ruff.

The Structural folds of Schwalbe are radial folds extending from the border of the ciliary and pupillary zones that are much broader and more widely-spaced, continuous with the "valleys" between the ciliary processes.

Some of the Circular contraction folds are a fine series of ridges that run near the pupillary margin and vary in thickness of the iris pigment epithelium; others are in ciliary portion of iris.[2] It changes colors like a rainbow.[citation needed]

Optic nerve
From Wikipedia, the free encyclopedia

This article is about the anatomical structure. For the comic book series, see Optic Nerve (comic). For the album about David Wojnarowicz, see Optic Nerve (CD-ROM).

This article includes a list of references, but its sources remain unclear because it has insufficient inline citations. Please help toimprove this article by introducing more precise citations. (November 2011)

Nerve: Optic Nerve

The left optic nerve and the optic tracts.

Latin

nervus opticus

Gray's

subject #197 882

MeSH

Optic+Nerve

The optic nerve, also known as cranial nerve II, transmits visual information from the retina to the brain. Derived from the embryonic retinal ganglion cell, a diverticulum located in the diencephalon, the optic nerve does not regenerate after transection.
Contents
[hide]

1 Anatomy 2 Physiology 3 Functional components 4 Role in disease 5 Additional images 6 See also 7 References

8 External links

Anatomy[edit]
The optic nerve is the second of twelve paired cranial nerves but is considered to be part of the central nervous system, as it is derived from an outpouching of the diencephalon during embryonic development. As a consequence, the fibers are covered with myelin produced byoligodendrocytes, rather than Schwann cells, which are found in the peripheral nervous system, and are encased within the meninges. Peripheral neuropathies like Guillain-Barr syndrome do not affect the optic nerve. The optic nerve is ensheathed in all three meningeal layers (dura, arachnoid, and pia mater) rather than the epineurium, perineurium, andendoneurium found in peripheral nerves. Fiber tracks of the mammalian central nervous system (as opposed to the peripheral nervous system) are incapable of regeneration, and, hence, optic nerve damage produces irreversible blindness. The fibres from the retina run along the optic nerve to nine primary visual nuclei in the brain, from which a major relay inputs into the primary visual cortex. The optic nerve is composed of retinal ganglion cell axons and support cells. It leaves the orbit (eye socket) via the optic canal, running postero-medially towards the optic chiasm, where there is a partial decussation (crossing) of fibres from the temporal visual fields (the nasal hemi-retina) of both eyes. The proportion of decussating fibers varies between species, and is correlated with the degree of binocular vision enjoyed by a species.[1] Most of the axons of the optic nerve terminate in the lateral geniculate nucleus from where information is relayed to the visual cortex, while other axons terminate in the pretectal nucleus and are involved in reflexive eye movements. Other axons terminate in the suprachiasmatic nucleus and are involved in regulating the sleep-wake cycle. Its diameter increases from about 1.6 mm within the eye to 3.5 mm in the orbit to 4.5 mm within the cranial space. The optic nerve component lengths are 1 mm in the globe, 24 mm in the orbit, 9 mm in the optic canal, and 16 mm in the cranial space before joining the optic chiasm. There, partial decussation occurs, and about 53% of the fibers cross to form the optic tracts. Most of these fibres terminate in the lateral geniculate body.[citation needed] From the lateral geniculate body, fibers of the optic radiation pass to the visual cortex in the occipital lobe of the brain. In more specific terms, fibers carrying information from the contralateral superior visual field traverse Meyer's loop to terminate in the lingual gyrus below the calcarine fissure in the occipital lobe, and fibers carrying information from the contralateral inferior visual field terminate more superiorly, to the cuneus.

Vitreous humour
From Wikipedia, the free encyclopedia

"Vitreous humor" redirects here. For the band, see Vitreous Humor.

Vitreous humour

Schematic diagram of the human eye.

Latin

humor vitreus

The vitreous humour (UK spelling) or vitreous humor (US spelling) is the clear gel that fills the space between the lens and the retina of theeyeball of humans and other vertebrates. It is often referred to as the vitreous body or simply "the vitreous".
Contents
[hide]

1 Composition, properties and function 2 Pathology 3 Clinical significance 4 Additional Images 5 See also 6 References 7 External links

Composition, properties and function[edit]

The vitreous is the transparent, colourless, gelatinous mass that fills the space between the lens of the eye and the retina lining the back of the eye. It is present at birth and does not change much over the course of aging. [1] It is produced by cells in the non-pigmented portion of the ciliary body deriven from embryonic mesenchyme cells which then degenerate after birth. [2] Its composition is similar to the cornea, but contains

very few cells (mostly phagocytes which remove unwanted cellular debris in the visual field, as well as the hyalocytes of the surface of the vitreous, which reprocess the hyaluronic acid), no blood vessels, and 98-99% of its volume is water (as opposed to 75% in the cornea) with salts, sugars, vitrosin (a type of collagen), a network of collagen type II fibres with the glycosaminoglycan hyaluronic acid, and also a wide array of proteins in micro amounts. Amazingly, with so little solid matter, it tautly holds the eye. The lens, on the other hand, is tightly packed with cells.[3] However, the vitreous has a viscosity two to four times that of pure water, giving it a gelatinous consistency. It also has a refractive index of 1.336.[4] Although the vitreous is in contact with the retina and helps to keep it in place by pressing it against the choroid, it does not adhere to the retina, except at the optic nerve disc, AKA: papilla nervi optici (where the retina sends about 1.2 million nerve fibres (axons) to the brain). It is also connected to the Ora serrata (where the retina ends anteriorly), at the Wieger-band, the dorsal side of the lens. It is however, not connected at the macula, the tiny spot in the retina which gives us our "detail" and central vision. Unlike the fluid in the frontal parts of the eye (aqueous humour) which is continuously replenished, the gel in the vitreous chamber is stagnant. Therefore, if blood, cells or other byproducts of inflammation get into the vitreous, they will remain there unless removed surgically[citation needed] (see floaters). If the vitreous pulls away from the retina, it is known as a vitreous detachment. As we age, the vitreous often liquefies and may collapse. This is more likely to occur, and occurs much earlier, in eyes that are nearsighted (myopia). It can also occur after injuries to the eye or inflammation in the eye (uveitis).

Pathology[edit]
The collagen fibres of the vitreous are held apart by electrical charges. With aging, these charges tend to reduce, and the fibres may clump together. Similarly, the gel may liquefy, a condition known as synaeresis, allowing cells and other organic clusters to float freely within the vitreous humour. These allow floaters which are perceived in the visual field as spots or fibrous strands. Floaters are generally harmless, but the sudden onset of recurring floaters may signify a posterior vitreous detachment (PVD) or other diseases of the eye.

Clinical significance[edit]
The metabolic exchange and equilibration between systemic circulation and vitreous humour is so slow that vitreous humour is sometimes the fluid of choice for postmortem analysis of glucoselevels or substances which would be more rapidly diffused, degraded, excreted, or metabolized from the general circulation.

Surface Ectoderm[edit]

The optic vesicles come into contact with the surface ectoderm which thickens to form the lens placode. The lens differentiates and invaginates until it pinches off from the ectoderm. The lens acts as an inducer back to the optic vesicle to transform it into the optic cup and back to the ectoderm to transform it into the cornea. The optic cup then delaminates into two layers: The neural retina and the retinal pigment epithelium. Surface ectoderm produces the following parts:

Lens corneal epithelium

Lens placode
From Wikipedia, the free encyclopedia Lens placode

Transverse section of head of chick embryo of fifty-two hours incubation.

Latin Precursor Gives rise to Code

placoda lentis surface ectoderm lens TE E5.16.3.1.0.0.13

The Lens placode is a thickened portion of ectoderm which serves as the precursor to the lens. SOX2 and Pou2f1 are involved in its development.[1]

Retinal pigment epithelium

From Wikipedia, the free encyclopedia

Retinal pigment epithelium

Section of retina. (Pigmented layer labeled at bottom right.)

Plan of retinal neurons. (Pigmented layer labeled at bottom right.)

Latin

Stratum pigmentosum retinae, pars pigmentosa retinae

Gray's

subject #225 1016

The pigmented layer of retina or retinal pigment epithelium (RPE) is the pigmented cell layer just outside the neurosensory retina that nourishes retinal visual cells, and is firmly attached to the underlying choroid and overlying retinal visual cells.[1][2]

Contents
[hide]

1 History 2 Anatomy 3 Function 4 Pathology 5 See also 6 References 7 External links

History[edit]

Choroid dissected from a calf's eye, showing black RPE and iridescent blue tapetum lucidum

The RPE was known in the 18th and 19th centuries as the pigmentum nigrum, referring to the observation that the RPE is dark (black in many animals, brown in humans); and as the tapetum nigrum, referring to the observation that in animals with a tapetum lucidum, in the region of the tapetum lucidum the RPE is not pigmented.[3]

Anatomy[edit]
The RPE is composed of a single layer of hexagonal cells that are densely packed with pigment granules.[1] At the ora serrata, the RPE continues as a membrane passing over the ciliary body and continuing as the back surface of the iris. This generates the fibers of the dilator. Directly beneath this epithelium is the neuroepithelium (i.e., rods and cones)passes jointly with the RPE. Both, combined, are understood to be the

ciliary epithelium of the embryo. The front end continuation of the retina is the posterior iris epithelium, which takes on pigment when it enters the iris[4] When viewed from the outer surface, these cells are smooth and hexagonal in shape. When seen in section, each cell consists of an outer non-pigmented part containing a large oval nucleus and an inner pigmented portion which extends as a series of straight thread-like processes between the rods, this being especially the case when the eye is exposed to light.

Function[edit]
The RPE shields the retina from excess incoming light. It supplies omega-3 fatty acids and glucose, the former for building photoreceptive membranes, the latter for energy. Retinal is supplied by the visual vitamin A cycle. Water is removed from the retinal side to the choroid side, at a rate of 1.4-11 microliters per square centimeter per hour. It maintains balance of pH, and routinely phagocytoses the oldest outer segment discs of the photoreceptors.[5] It has a self-contained immune system, which is connected with the immune system proper to either shut down interactions when healthy, and when there is disease, it teams up with the main immune controls. Finally, it secretes substances to help build and sustain the choroid and retina. [6] The retinal pigment epithelium also serves as the limiting transport factor that maintains the retinal environment by supplying small molecules such as amino acid, ascorbic acid and D-glucose while remaining a tight barrier to choroidal blood borne substances. Homeostasis of the ionic environment is maintained by a delicate transport exchange system. In some clinical studies, RPE auto transplant has been used in treating Macular degeneration. Also experimental studies have reported in vitro expanded RPE used in similar studies.[7]

Pathology[edit]
In the eyes of albinos, the cells of this layer contain no pigment. Dysfunction of the RPE is found in AgeRelated Macular Degeneration and Retinitis Pigmentosa.

Neural crest[edit]

Neural crest cells are themselves derived from the ectoderm and lie close to the neural tube:

Sclera Corneal stroma and endothelium Connective tissue and bony structure of the orbit

Neural crest
From Wikipedia, the free encyclopedia Neural crest

The formation of neural crest during the process of neurulation. Neural crest is first induced in the region of the neural plate border. After neural tube closure, neural crest delaminates from the region between the dorsal neural tube and overlying ectoderm and migrates out towards the periphery.

Latin Code

crista neuralis TE E5.0.2.1.0.0.2

Neural crest cells are a transient, multipotent, migratory cell population unique to vertebrates that gives rise to a diverse cell lineage includingmelanocytes, craniofacial cartilage and bone, smooth muscle, peripheral and enteric neurons and glia.[1]

After gastrulation, neural crest cells are specified at the border of the neural plate and the nonneural ectoderm. During neurulation, the borders of the neural plate, also known as the neural folds,

converge at the dorsal midline to form the neural tube. Subsequently, neural crest cells from the roof plate of the neural tube undergo an epithelial to mesenchymal transition, delaminating from the neuroepithelium and migrating through the periphery where they differentiate into varied cell types.[1] The emergence of neural crest was important in vertebrate evolution because many of its structural derivatives are defining features of the vertebrate clade.[2]

Underlying the development of neural crest is a gene regulatory network, described as a set of interacting signals, transcription factors, and downstream effector genes that confer cell characteristics such as multipotency and migratory capabilities.[3] Understanding the molecular mechanisms of neural crest formation is important for our knowledge of human disease because of its contributions to multiple cell lineages. Abnormalities in neural crest development cause neurocristopathies, which include conditions such as frontonasal dysplasia, Waardenburg-Shah syndrome, and DiGeorge syndrome.[1]

Therefore, defining the mechanisms of neural crest development may reveal key insights into vertebrate evolution and neurocristopathies.

Sclera
From Wikipedia, the free encyclopedia

"White of the eye" redirects here. For the film, see White of the Eye (film).

Sclera

The sclera, as separated from the cornea by the corneal limbus.

Gray's subject #225 1006

Artery anterior ciliary arteries, long posterior ciliary arteries,short posterior ciliary arteries

MeSH Sclera

The sclera (from the Greek skleros, meaning hard[1]), also known as the white of the eye, is the opaque, fibrous, protective, outer layer of the eye containing collagen and elastic fiber.[2] In humans the whole sclera is white, contrasting with the coloured iris, but in other mammals the visible part of the sclera matches the colour of the iris, so the white part does not normally show. In the development of theembryo, the sclera is derived from the neural crest.[3] In children, it is thinner and shows some of the underlying pigment, appearing slightly blue. In the elderly, fatty deposits on the sclera can make it appear slightly yellow.

The human eye is relatively rare for having an iris which is small enough for its position to be plainly visible against the sclera. This makes it easier for one individual to infer where another individual is looking, and it may have evolved as a method of nonverbal communication.

Cornea
From Wikipedia, the free encyclopedia

For other uses, see Cornea (disambiguation).

Cornea

Schematic diagram of the human eye showing the cornea as separated from the sclera by the corneal limbus

Vertical section of human cornea from near the margin. (Waldeyer.) Magnified.

1. 2. 3. 4. 5.

Epithelium. Anterior elastic lamina. substantia propria. Posterior elastic lamina (Descemet's membrane). Endothelium of the anterior chamber. 1. Oblique fibers in the anterior layer of the substantia propria. 2. Lamell the fibers of which are cut across, producing a dotted appearance. 3. Corneal corpuscles appearingfusiform in section. d. Lamell the fibers of which are cut longitudinally. 4. Transition to the sclera, with more distinct fibrillation, and surmounted by a thicker epithelium. 5. Small bloodvessels cut across near the margin of the cornea.

Gray's

subject #225 1070

The cornea is the transparent front part of the eye that covers the iris, pupil, and anterior chamber. The cornea, with the anterior chamber and lens,refracts light, with the cornea accounting for approximately two-thirds of the eye's total optical power.[1][2] In humans, the refractive power of the cornea is approximately 43 dioptres.[3] While the cornea contributes most of the eye's focusing power, its focus is fixed. The curvature of the lens, on the other hand, can be adjusted to "tune" the focus depending upon the object's distance. Medical terms related to the cornea often start with the prefix "kerat-" from the Greek word , horn.

Mesoderm[edit] Mesoderm contributes to the following structures:

Extraocular muscles Endothelial lining of blood vessels of the eye

Developmental cascade[edit]
According to Liem et al., the organogenesis of the eye is pointed out as an example of a developmental cascade of inductions. The eye is essentially a derivative of the ectoderm from the somatic ectoderm and neural tube, with a succession of inductions by the chordamesoderm. Chordamesoderm induces the anterior portion of the neural tube to form the precursors of the synapomorphic tripartite brain of vertebrates, and it will form a bulge called the diencephalon. Further induction by the chordamesoderm will form a protrusion: the optic visicle. This visicle will be subsequently invaginated by means of further inductions from the chordamesoderm. The optic visicle will then induce the ectoderm that thickens (lens placode) and further invaginates to a point that detaches from the ectoderm and forms a neurogenic placode by itself. The lens placode is affected by the chordamesoderm making it to invaginate and forms the optic cup composed by an outer layer of neural retina and inner layer the pigmented retina that will unite and form the optic stalk. The pigmented retina is formed by rods and cones and composed of small cilia typical of the ependymal epithelium of the neural tube. Some cells in the lens vesicle will be fated to form the cornea and the lens vesicle will develop completely to form the definitive lens. Iris is formed from the optic cup cells.

Responsivity of head epidermis[edit]

Only the epidermis in the head is competent to respond to the signal from the optic vesicles. Both the optic vesicle and the head epidermis are required for eye development. The competence of the head epidermis to respond to the optic vesicle signals comes from the expression of Pax6 in the epidermis. Pax6 is necessary and sufficient for eye induction. This competenece is acquired gradually during gastrulation and neurulation from interactions with the endoderm, mesoderm, and neural plate.

Regulation and inhibition[edit]

Sonic hedgehog reduces the expression of Pax6. When Shh is inhibited during development, the domain of expression for Pax6 is expanded and the eyes fail to separate causing cyclopia.[3] Overexpression of Shh causes a loss of eye structures.

Retinoic acid generated from vitamin A in the retina plays an essential role in eye development as a secreted paracrine signal which restricts invasion of perioptic mesenchyme around the optic cup.[4] Vitamin A deficiencyduring embryogenesis results in anterior segment defects (particularly cornea and eyelids) that lead to vision loss or blindness. There is some evidence that LMX1B plays a role in periocular mesenchymal survival.[5]