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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 (anatomy), iris of the Human eye, eye that allows light to strike the retina.Cassin, B. and Solomon, S. (1990) ''Dictionary of Eye Terminology''. Gainesville, Florida: Triad Publishing ...
, 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 Eye color is a polygenic phenotypic character determined by two distinct factors: the pigmentation of the eye's iris and the frequency-dependence of the scattering of light by the turbid medium in the stroma of the iris. In humans, the ...
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
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 compounds whereas pigments are often inorganic compou ...
ed fibrovascular layer known as a stroma and, beneath the stroma, pigmented epithelial cells. The stroma is connected to a
sphincter A sphincter is a circular muscle that normally maintains constriction of a natural body passage or orifice and which relaxes as required by normal physiological functioning. Sphincters are found in many animals. There are over 60 types in the h ...
muscle ( sphincter pupillae), which contracts the pupil in a circular motion, and a set of dilator muscles (
dilator pupillae The iris dilator muscle (pupil dilator muscle, pupillary dilator, radial muscle of iris, radiating fibers), is a smooth muscle of the eye, running radially in the iris and therefore fit as a dilator. The pupillary dilator consists of a spokelike ...
), 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 An encyclopedia (American English) or encyclopædia (British English) is a reference work or compendium providing summaries of knowledge either general or special to a particular field or discipline. Encyclopedias are divided into articles ...
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 containing mainly collagen and some crucial elastic fiber. In humans, and so ...
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 The trabecular meshwork is an area of tissue in the eye located around the base of the cornea, near the ciliary body, and is responsible for draining the aqueous humor from the eye via the anterior chamber (the chamber on the front of the ...
, 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 Intraocular pressure (IOP) is the fluid pressure inside the eye. 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 tonometers are calibrated t ...
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, so-called because it has no sarcomeres and therefore no striations (''bands'' or ''stripes''). It is divided into two subgroups, single-unit and multiunit smooth muscle. Within single-unit ...
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, in An ...
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 The stroma of the iris is a fibrovascular layer of tissue. It is the upper layer of two in the iris. Structure The stroma is a delicate interlacement of fibres. Some circle the circumference of the iris and the majority radiate toward the pupi ...
*
Iris sphincter muscle The iris sphincter muscle (pupillary sphincter, pupillary constrictor, circular muscle of iris, circular fibers) is a muscle in the part of the eye called the iris. It encircles the pupil of the iris, appropriate to its function as a constricto ...
*
Iris dilator muscle The iris dilator muscle (pupil dilator muscle, pupillary dilator, radial muscle of iris, radiating fibers), is a smooth muscle of the eye, running radially in the iris and therefore fit as a dilator. The pupillary dilator consists of a spokelike ...
(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 vertebrates that arise from the embryonic ectoderm germ layer, and in turn give rise to a diverse cell lineage—including melanocytes, craniofacial cartilage and bone, smooth muscle, per ...
, 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 compounds whereas pigments are often inorganic compou ...
ed, 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 Albinism is the congenital absence of melanin in an animal or plant resulting in white hair, feathers, scales and skin and pink or blue eyes. Individuals with the condition are referred to as albino. Varied use and interpretation of the term ...
, 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 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 and fi ...
. 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 optical ...
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 containing mainly collagen and some crucial elastic fiber. In humans, and so ...
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 Heterochromia is a variation in coloration. The term is most often used to describe color differences of the iris, but can also be applied to color variation of hair or skin. Heterochromia is determined by the production, delivery, and concent ...
, below). Colors other than brown or black are due to selective reflection and absorption from the other stromal components. Sometimes,
lipofuscin Lipofuscin is the name given to fine yellow-brown pigment granules composed of lipid-containing residues of lysosomal digestion. It is considered to be one of the aging or "wear-and-tear" pigments, found in the liver, kidney, heart muscle, reti ...
, 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 Collagen () is the main structural protein in the extracellular matrix found in the body's various connective tissues. As the main component of connective tissue, it is the most abundant protein in mammals, making up from 25% to 35% of the whol ...
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 of th ...
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 Diffraction is defined as the interference or bending of waves around the corners of an obstacle or through an aperture into the region of geometrical shadow of the obstacle/aperture. The diffracting object or aperture effectively becomes a s ...
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 ...
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 Waardenburg syndrome is a group of rare genetic conditions characterised by at least some degree of congenital hearing loss and pigmentation deficiencies, which can include bright blue eyes (or one blue eye and one brown eye), a white foreloc ...
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 Shepherd The Australian Shepherd is a breed of herding dog from the United States. The name of the breed is technically a misnomer, as it was developed in California in the 19th century; the breed was unknown in Australia at the time. It is claimed that A ...
s and
Border Collie The Border Collie is a Scottish breed of herding dog of medium size. Widely considered to be the most intelligent dog breed, they are descended from landrace sheepdogs once found all over the British Isles, but became standardised in the Ang ...
s) 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 Glaucoma is a group of eye diseases that result in damage to the optic nerve (or retina) and cause vision loss. The most common type is open-angle (wide angle, chronic simple) glaucoma, in which the drainage angle for fluid within the eye rem ...
*
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 Uveitis () is inflammation of the uvea, the pigmented layer of the eye between the inner retina and the outer fibrous layer composed of the sclera and cornea. The uvea consists of the middle layer of pigmented vascular structures of the eye and ...
*
Iritis Uveitis () is inflammation of the uvea, the pigmented layer of the eye between the inner retina and the outer fibrous layer composed of the sclera and cornea. The uvea consists of the middle layer of pigmented vascular structures of the eye and ...
* Miosis/
Mydriasis Mydriasis is the dilation of the pupil, usually having a non-physiological cause, or sometimes a physiological pupillary response. Non-physiological causes of mydriasis include disease, trauma, or the use of certain types of drugs. Normally, a ...
* 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 despite lacking biological plausibility, testability, repeatability, or evidence from clinical trials. Complementary medicine (CM), complementary and al ...
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 A rainbow is a meteorological phenomenon that is caused by reflection, refraction and dispersion of light in water droplets resulting in a spectrum of light appearing in the sky. It takes the form of a multicoloured circular arc. Rainbows c ...
", 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 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 presen ...
*
Gonioscopy Gonioscopy a routine ophthalmological procedure that measures the angle between the iris and the cornea (the iridocorneal angle), using a goniolens (also known as a gonioscope) together with a slit lamp or operating microscope. Its use is import ...
*
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 Iridoplegia is the paralysis of the sphincter of the iris. It can occur in due to direct orbital injury, which may result in short lived blurred vision. Types It can be of three types: # accommodative iridoplegia- Noncontraction of pupils during ...
– paralysis of the sphincter muscles of the iris *
Iris recognition Iris recognition is an automated method of biometric identification that uses mathematical pattern-recognition techniques on video images of one or both of the irises of an individual's eyes, whose complex patterns are unique, stable, and can ...
*
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