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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 The pupil is a black hole located in the center of the iris of the eye that allows light to strike the retina The retina (from la, rete "net") is the innermost, light-sensitive layer of tissue of the eye of most vertebrate Vertebr ...
, 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 vertebrate Vertebrates () comprise all animal Animals are multicellular, eukaryotic organisms in the biological kingdom An ...
. 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. A ...
, while the iris is the diaphragm.


Structure

The iris consists of two layers: the front
pigment A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at some stage in their use. Generally dyes are often organic compound In chemistry, organic compounds are gen ...
ed 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 Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, ...
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 The sclera, also known as the white of the eye or, in older literature, as the tunica albuginea oculi, is the opaque, fibrous, protective, outer layer of the human eye The human eye is a sensory organ, part of the sensory nervous system, th ...
and the anterior ciliary body. The iris and ciliary body together are known as the anterior
uvea The uvea (; Lat. ''uva'', "grape"), also called the ''uveal layer'', ''uveal coat'', ''uveal tract'', ''vascular tunic'' or ''vascular layer'' is the pigmented middle of the three concentric layers that make up an eye. History and etymolog ...
. 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 Intraocular pressure (IOP) is the fluid pressure inside the human eye, eye. Ocular tonometry, Tonometry is the method eye care professionals use to determine this. IOP is an important aspect in the evaluation of patients at risk of glaucoma. Most ...
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 Smooth muscle is an involuntary non-striated muscle Striations means a series of ridges, furrows or linear marks, and is used in several ways: * Glacial striation * Striation (fatigue), in material * Striation (geology), a ''striation'' as ...
in mammals and amphibians, but are
striated muscle Striations means a series of ridges, furrows or linear marks, and is used in several ways: * Glacial striation * Striation (fatigue), in material * Striation (geology), a ''striation'' as a result of a geological fault * Striation Valley, i ...
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 Neural crest cells are a temporary group of cells unique to vertebrate Vertebrates () comprise all animal Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume or ...
, 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
pigment A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at some stage in their use. Generally dyes are often organic compound In chemistry, organic compounds are gen ...
ed, with the
color Color (American English American English, sometimes called United States English or U.S. English, is the set of varieties of the English language English is a West Germanic language of the Indo-European language family, with ...
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 pigment A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at som ...
. 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 Vertebrates () comprise all animal Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exception ...
and
hair Hair is a protein filament that grows from follicles found in the dermis. Hair is one of the defining characteristics of mammals. The human body, apart from areas of glabrous skin, is covered in follicles which produce thick terminal an ...
. 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 In biology, epigenetics is the study of stable phenotypic changes (known as ''marks'') that do not involve alterations in the DNA sequence. The Greek prefix '' epi-'' ( "over, outside of, around") in ''epigenetics'' implies features that are ...
constitution in this context. A person's "eye color" is actually the color of one's iris, the
cornea The cornea is the transparent front part of the eye that covers the iris, pupil, and anterior chamber. Along with the anterior chamber and lens, the cornea refracts light, accounting for approximately two-thirds of the eye's total optica ...
being transparent and the white
sclera The sclera, also known as the white of the eye or, in older literature, as the tunica albuginea oculi, is the opaque, fibrous, protective, outer layer of the human eye The human eye is a sensory organ, part of the sensory nervous system, th ...
entirely outside the area of interest. Melanin is yellowish to dark hazel in the stromal pigment cells, and black in the iris pigment epithelium, 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, 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 Iron () is a chemical element A chemical element is a species of atom ...
in the blood vessels,
collagen Collagen () is the main structural protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalys ...
in the vessel and stroma) is the most important element.
Rayleigh scattering Rayleigh scattering ( ), named after the 19th-century British physicist Lord Rayleigh (John William Strutt), is the predominantly elastic scattering of light or other electromagnetic radiation by particles much smaller than the wavelength ...
and Tyndall scattering, (which also happen in the sky) and diffraction also occur. Raman scattering, 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
wing A wing is a type of fin that produces lift while moving through air or some other fluid. Accordingly, wings have streamlined cross-sections that are subject to aerodynamic forces and act as airfoils. A wing's aerodynamic efficiency is e ...
s, 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 *
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 lesio ...
*
Iridocyclitis Uveitis () is inflammation of the uvea, the pigmented layer of the eye between the inner retina The retina (from la, rete "net") is the innermost, light-sensitive layer of tissue of the eye of most vertebrate Vertebrates () comprise ...
* Iritis *
Miosis Miosis, or myosis (), is excessive constriction of the pupil.Farlex medical dictionary
citing: ...
/
Mydriasis Mydriasis is the dilation of the pupil The pupil is a black hole located in the center of the iris of the eye that allows light to strike the retina The retina (from la, rete "net") is the innermost, light-sensitive layer of tissue ...
* Synechia * Third nerve palsy *
Aniridia Aniridia is the absence of the iris, a muscular structure that opens and closes the pupil to allow light into the eye. It is also responsible for eye color. Without it, the central eye appears all black. It can be congenital, in which both ey ...


Alternative medicine


Iridology

Iridology (also known as iridodiagnosis) is an
alternative medicine Alternative medicine is any practice that aims to achieve the healing effects of medicine Medicine is the science Science is a systematic endeavor that builds and organizes knowledge in the form of testable explanations and pr ...
technique whose proponents believe that patterns, colors, and other characteristics of the iris can be examined to determine information about a patient's systemic 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 A rainbow is a meteorological Meteorology is a branch of the atmospheric sciences (which include atmospheric chemistry and physics) with a major focus on weather forecasting. The study of meteorology dates back millennia, though signi ...
", 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 Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world Ancient history is a ...
'', because of the many colours 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 Aniridia is the absence of the iris, a muscular structure that opens and closes the pupil to allow light into the eye. It is also responsible for eye color. Without it, the central eye appears all black. It can be congenital, in which both ey ...
* Blood–ocular barrier *
Coloboma A coloboma (from the Greek , meaning defect) is a hole in one of the structures of the eye, such as the iris, retina, choroid, or optic disc. The hole is present from birth and can be caused when a gap called the choroid fissure, which is p ...
* Gonioscopy *
Human eye The human eye is a sensory organ, part of the sensory nervous system, that reacts to visible light and allows humans to use visual information for various purposes including seeing things, keeping balance, and maintaining circadian rhythm. ...
* 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 vertebrate Vertebrates () comprise all animal Animals are multicellular, eukaryotic organisms in the biological kingdom An ...


References


External links


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