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In humans and most mammals and birds, the iris (plural: ''irides'' or ''irises'') is a thin, annular structure in the eye, responsible for controlling the diameter and size of the pupil, and thus the amount of light reaching the
retina The retina (from la, rete "net") is the innermost, light-sensitive layer of tissue of the eye of most vertebrates and some molluscs. The optics of the eye create a focused two-dimensional image of the visual world on the retina, which the ...
. Eye color is defined by the iris. In optical terms, the pupil is the eye's
aperture In optics, an aperture is a hole or an opening through which light travels. More specifically, the aperture and focal length of an optical system determine the cone angle of a bundle of rays that come to a focus in the image plane. An ...
, while the iris is the
diaphragm Diaphragm may refer to: Anatomy * Thoracic diaphragm, a thin sheet of muscle between the thorax and the abdomen * Pelvic diaphragm or pelvic floor, a pelvic structure * Urogenital diaphragm or triangular ligament, a pelvic structure Other * Diap ...
.


Structure

The iris consists of two layers: the front pigmented fibrovascular layer known as a stroma and, beneath the stroma, pigmented epithelial cells. The stroma is connected 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 sphincter pupillae is the opposing muscle of the dilator pupillae. The pupil's diameter, and thus the inner border of the iris, changes size when constricting or dilating. The outer border of the iris does not change size. The constricting muscle is located on the inner border. The back surface is covered by a heavily pigmented
epithelial Epithelium or epithelial tissue is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. It is a thin, continuous, protective layer of compactly packed cells with a little intercellu ...
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."eye, human." Encyclopædia Britannica from Encyclopædia Britannica 2006 Ultimate Reference Suite DVD The outer edge of the iris, known as the root, is attached to the sclera and the anterior
ciliary body The ciliary body is a part of the eye that includes the ciliary muscle, which controls the shape of the lens, and the ciliary epithelium, which produces the aqueous humor. The aqueous humor is produced in the non-pigmented portion of the ciliar ...
. 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 The aqueous humour is a transparent water-like fluid similar to plasma, but containing low protein concentrations. It is secreted from the ciliary body, a structure supporting the lens of the eyeball. It fills both the anterior and the posteri ...
constantly drains out of the eye, with the result that diseases of the iris often have important effects on intraocular pressure and indirectly on vision. The iris along with the anterior ciliary body provide a secondary pathway for aqueous humour to drain from the eye. The iris is divided into two major regions: # The pupillary zone is the inner region whose edge forms the boundary of the pupil. # 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 vestige of the coating of the embryonic pupil. 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.Gold, Daniel H; Lewis, Richard; "Clinical Eye Atlas," pp. 396–397 The muscle cells of the iris are smooth muscle in mammals and amphibians, but are striated muscle in reptiles (including birds). Many fish have neither, and, as a result, their irises are unable to dilate and contract, so that the pupil always remains of a fixed size.


Front

*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 irises. *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. *Crypts on the base of the iris are additional openings that can be observed close to the outermost part of the ciliary portion of the iris.


Back

*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.


Microanatomy

From anterior (front) to posterior (back), the layers of the iris are: *Anterior limiting layer * Stroma of iris * Iris sphincter muscle * Iris dilator muscle (myoepithelium) *Anterior pigment epithelium *Posterior pigment epithelium


Development

The stroma and the anterior border layer of the iris are derived from the neural crest, and behind the stroma of the iris, the sphincter pupillae and dilator pupillae muscles, as well as the iris epithelium, develop from optic cup neuroectoderm.


Eye color

The iris is usually strongly pigmented, with the
color Color (American English) or colour (British English) is the visual perceptual property deriving from the spectrum of light interacting with the photoreceptor cells of the eyes. Color categories and physical specifications of color are associ ...
typically ranging between brown, hazel, green, gray, and blue. Occasionally, the color of the iris is due to a lack of pigmentation, as in the pinkish-white of oculocutaneous albinism, or to obscuration of its pigment by blood vessels, as in the red of an abnormally vascularised iris. Despite the wide range of colors, the only pigment that contributes substantially to normal human iris color is the dark pigment
melanin Melanin (; from el, μέλας, melas, black, dark) is a broad term for a group of natural pigments found in most organisms. Eumelanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the amino ...
. The quantity of melanin pigment in the iris is one factor in determining the phenotypic eye color of a person. Structurally, this huge molecule is only slightly different from its equivalent found in
skin Skin is the layer of usually soft, flexible outer tissue covering the body of a vertebrate animal, with three main functions: protection, regulation, and sensation. Other animal coverings, such as the arthropod exoskeleton, have different ...
and hair. Iris color is due to variable amounts of eumelanin (brown/black melanins) and pheomelanin (red/yellow melanins) produced by melanocytes. More of the former is found in brown-eyed people and of the latter in blue- and green-eyed people.


Genetic and physical factors determining iris color

Iris color is a highly complex phenomenon consisting of the combined effects of texture, pigmentation, fibrous tissue, and blood vessels within the iris stroma, which together make up an individual's epigenetic constitution in this context. A person's "eye color" is actually the color of one's iris, the cornea being transparent and the white sclera entirely outside the area of interest. Melanin is yellowish to dark hazel in the stromal pigment cells, and black in the
iris pigment epithelium The iris pigment epithelium (IPE) is a one cell thick layer of cuboidal cells lying behind the iris. The epithelial cells are highly pigmented due to the numerous large melanosomes which pack the cytoplasm of each cell. Towards the central axis, ...
, which lies in a thin but very opaque layer across the back of the iris. Most human irises also show a condensation of the brownish stromal melanin in the thin anterior border layer, which by its position has an overt influence on the overall color. The degree of dispersion of the melanin, which is in subcellular bundles called
melanosomes A melanosome is an organelle found in animal cells and is the site for synthesis, storage and transport of melanin, the most common light-absorbing pigment found in the animal kingdom. Melanosomes are responsible for color and photoprotection ...
, has some influence on the observed color, but melanosomes in the iris of humans and other vertebrates are not mobile, and the degree of pigment dispersion cannot be reversed. Abnormal clumping of melanosomes does occur in disease and may lead to irreversible changes in iris color (see heterochromia, below). Colors other than brown or black are due to selective reflection and absorption from the other stromal components. Sometimes, lipofuscin, a yellow "wear and tear" pigment, also enters into the visible eye color, especially in aged or diseased green eyes. The optical mechanisms by which the nonpigmented stromal components influence eye color are complex, and many erroneous statements exist in the literature. Simple selective absorption and reflection by biological molecules (
hemoglobin Hemoglobin (haemoglobin BrE) (from the Greek word αἷμα, ''haîma'' 'blood' + Latin ''globus'' 'ball, sphere' + ''-in'') (), abbreviated Hb or Hgb, is the iron-containing oxygen-transport metalloprotein present in red blood cells (erythroc ...
in the blood vessels, collagen in the vessel and stroma) is the most important element. Rayleigh scattering and
Tyndall scattering The Tyndall effect is light scattering by particles in a colloid or in a very fine suspension. Also known as Tyndall scattering, it is similar to Rayleigh scattering, in that the intensity of the scattered light is inversely proportional to the ...
, (which also happen in the sky) and diffraction also occur.
Raman scattering Raman scattering or the Raman effect () is the inelastic scattering of photons by matter, meaning that there is both an exchange of energy and a change in the light's direction. Typically this effect involves vibrational energy being gained by ...
, and constructive interference, as in the feathers of birds, do not contribute to the color of the human eye, but interference phenomena are important in the brilliantly colored iris pigment cells ( iridophores) in many animals. Interference effects can occur at both molecular and light-microscopic scales, and are often associated (in melanin-bearing cells) with quasicrystalline formations, which enhance the optical effects. Interference is recognised by characteristic dependence of color on the angle of view, as seen in eyespots of some
butterfly Butterflies are insects in the macrolepidopteran clade Rhopalocera from the order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The group compris ...
wings, although the chemical components remain the same. White babies are usually born blue-eyed since no pigment is in the stroma, and their eyes appear blue due to scattering and selective absorption from the posterior epithelium. If melanin is deposited substantially, brown or black color is seen; if not, they will remain blue or gray."Sensory Reception: Human Vision: Structure and function of the Human Eye" vol. 27, p. 175 Encyclopædia Britannica, 1987 All the contributing factors towards eye color and its variation are not fully understood. Autosomal recessive/dominant traits in iris color are inherent in other species, but coloration can follow a different pattern.


Different colors in the two eyes

Heterochromia (also known as a heterochromia iridis or heterochromia iridum) is an ocular condition in which one iris is a different color from the other iris (complete heterochromia), or where the part of one iris is a different color from the remainder (partial heterochromia or sectoral heterochromia). Uncommon in humans, it is often an indicator of ocular disease, such as chronic iritis or diffuse iris melanoma, but may also occur as a normal variant. Sectors or patches of strikingly different colors in the same iris are less common. Anastasius the First was dubbed ''dikoros'' (having two irises) for his patent heterochromia since his right iris had a darker color than the left one. In contrast, heterochromia and variegated iris patterns are common in veterinary practice. Siberian Husky dogs show heterochromia, possibly analogous to the genetically determined Waardenburg syndrome of humans. Some white cat fancies (e.g., white Turkish Angora or white Turkish van cats) may show striking heterochromia, with the most common pattern being one uniformly blue, the other copper, orange, yellow, or green. Striking variation within the same iris is also common in some animals, and is the norm in some species. Several herding breeds, particularly those with a blue merle coat color (such as Australian Shepherds and Border Collies) may show well-defined blue areas within a brown iris, as well as separate blue and darker eyes. Some horses (usually within the white, spotted, palomino, or cremello groups of breeds) may show amber, brown, white and blue all within the same eye, without any sign of eye disease. One eye with a white or bluish-white iris is also known as a "walleye".


Clinical significance

* Angle closure glaucoma *
Anisocoria Anisocoria is a condition characterized by an unequal size of the eyes' pupils. Affecting up to 20% of the population, anisocoria is often entirely harmless, but can be a sign of more serious medical problems. Causes Anisocoria is a common condit ...
*
Horner's syndrome Horner's syndrome, also known as oculosympathetic paresis, is a combination of symptoms that arises when a group of nerves known as the sympathetic trunk is damaged. The signs and symptoms occur on the same side (ipsilateral) as it is a lesion ...
* Iridocyclitis * Iritis * Miosis/ Mydriasis * Synechia * Third nerve palsy * Aniridia


Alternative medicine


Iridology

Iridology (also known as iridodiagnosis) is an alternative medicine technique whose proponents believe that patterns, colors, and other characteristics of the iris can be examined to determine information about a patient's
systemic Systemic fundamental to a predominant social, economic, or political practice. This refers to: In medicine In medicine, ''systemic'' means affecting the whole body, or at least multiple organ systems. It is in contrast with ''topical'' or ''loc ...
health. Practitioners match their observations to "iris charts", which divide the iris into zones corresponding to specific parts of the human body. Iridologists see the eyes as "windows" into the body's state of health. Iridology is not supported by quality research studies, and is considered pseudoscience''Iridology Is Nonsense'', a web page with further references
/ref> by the majority of medical practitioners and eye-care professionals.


Etymology

The word "iris" is derived from the Greek word for " rainbow", also its goddess plus messenger of the gods in the ''
Iliad The ''Iliad'' (; grc, Ἰλιάς, Iliás, ; "a poem about Ilium") is one of two major ancient Greek epic poems attributed to Homer. It is one of the oldest extant works of literature still widely read by modern audiences. As with the ''Ody ...
'', because of the many
colours Color (American English) or colour (British English) is the visual perceptual property deriving from the spectrum of light interacting with the photoreceptor cells of the eyes. Color categories and physical specifications of color are associa ...
of this eye part.


Graphics

File:Three Main Layers of the Eye.png, Structures of the eye Image:Gray878.png, Iris, front view File:Fluorescein angiograpy of iris reveals radial layout of blood vessels.jpg, Fluorescein angiograpy of the iris reveals a radial layout of blood vessels.


See also

* Aniridia * Blood–ocular barrier * Coloboma * Gonioscopy * Human eye * Iridoplegia – paralysis of the sphincter muscles of the iris * Iris recognition *
Retina The retina (from la, rete "net") is the innermost, light-sensitive layer of tissue of the eye of most vertebrates and some molluscs. The optics of the eye create a focused two-dimensional image of the visual world on the retina, which the ...


References


External links


Detailed photographs of human irides
* * – "Sagittal Section Through the Eyeball" {{Authority control Eye color