Eye - Knowledge (XXG)

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whole visual field; this fast response is further accelerated by the large nerve bundles which rush the information to the brain. Focusing the image would also cause the sun's image to be focused on a few receptors, with the possibility of damage under the intense light; shielding the receptors would block out some light and thus reduce their sensitivity. This fast response has led to suggestions that the ocelli of insects are used mainly in flight, because they can be used to detect sudden changes in which way is up (because light, especially UV light which is absorbed by vegetation, usually comes from above).

1364:, or how much an eye can differentiate one object from another in terms of visual angles. Resolution in CPD can be measured by bar charts of different numbers of white/black stripe cycles. For example, if each pattern is 1.75 cm wide and is placed at 1 m distance from the eye, it will subtend an angle of 1 degree, so the number of white/black bar pairs on the pattern will be a measure of the cycles per degree of that pattern. The highest such number that the eye can resolve as stripes, or distinguish from a grey block, is then the measurement of visual acuity of the eye. 1557: 650:, are eye-spots which may be set into a pit to reduce the angles of light that enters and affects the eye-spot, to allow the organism to deduce the angle of incoming light. Found in about 85% of phyla, these basic forms were probably the precursors to more advanced types of "simple eyes". They are small, comprising up to about 100 cells covering about 100 μm. The directionality can be improved by reducing the size of the aperture, by incorporating a reflective layer behind the receptor cells, or by filling the pit with a refractile material. 1406:"Colour vision is the faculty of the organism to distinguish lights of different spectral qualities." All organisms are restricted to a small range of electromagnetic spectrum; this varies from creature to creature, but is mainly between wavelengths of 400 and 700 nm. This is a rather small section of the electromagnetic spectrum, probably reflecting the submarine evolution of the organ: water blocks out all but two small windows of the EM spectrum, and there has been no evolutionary pressure among land animals to broaden this range. 4077: 87: 1545: 806: 1413:, has a peak response at 500 nm. Small changes to the genes coding for this protein can tweak the peak response by a few nm; pigments in the lens can also filter incoming light, changing the peak response. Many organisms are unable to discriminate between colours, seeing instead in shades of grey; colour vision necessitates a range of pigment cells which are primarily sensitive to smaller ranges of the spectrum. In primates, geckos, and other organisms, these take the form of 297: 916: 908: 456: 1182: 308: 775: 1294:

eyes are constrained to a spherical shape, but other forms of compound eyes may deform to a shape where more ommatidia are aligned to, say, the horizon, without altering the size or density of individual ommatidia. Eyes of horizon-scanning organisms have stalks so they can be easily aligned to the horizon when this is inclined, for example, if the animal is on a slope.

1248:, thought to be the protovertebrate, were evidently pushed to very deep, dark waters, where they were less vulnerable to sighted predators, and where it is advantageous to have a convex eye-spot, which gathers more light than a flat or concave one. This would have led to a somewhat different evolutionary trajectory for the vertebrate eye than for other animal eyes. 69: 1108:. The shrimp has an eye of the refracting superposition type, in the rear behind this in each eye there is a single large facet that is three times in diameter the others in the eye and behind this is an enlarged crystalline cone. This projects an upright image on a specialised retina. The resulting eye is a mixture of a simple eye within a compound eye. 1476:. They require brighter light to function than rods require. In humans, there are three types of cones, maximally sensitive to long-wavelength, medium-wavelength, and short-wavelength light (often referred to as red, green, and blue, respectively, though the sensitivity peaks are not actually at these colours). The colour seen is the combined effect of 1004:

type of compound eye, for which a minimal size exists below which effective superposition cannot occur, is normally found in nocturnal insects, because it can create images up to 1000 times brighter than equivalent apposition eyes, though at the cost of reduced resolution. In the parabolic superposition compound eye type, seen in arthropods such as

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Prey animals and competing predators alike would be at a distinct disadvantage without such capabilities and would be less likely to survive and reproduce. Hence multiple eye types and subtypes developed in parallel (except those of groups, such as the vertebrates, that were only forced into the photopic environment at a late stage).

897: 794:, reptiles, and most other terrestrial vertebrates (along with spiders and some insect larvae) the vitreous fluid has a higher refractive index than the air. In general, the lens is not spherical. Spherical lenses produce spherical aberration. In refractive corneas, the lens tissue is corrected with inhomogeneous lens material (see 931:, singular). The image perceived is a combination of inputs from the numerous ommatidia (individual "eye units"), which are located on a convex surface, thus pointing in slightly different directions. Compared with simple eyes, compound eyes possess a very large view angle, and can detect fast movement and, in some cases, the 1256:, and allowed functionality outside of water. The transparent protective cells eventually split into two layers, with circulatory fluid in between that allowed wider viewing angles and greater imaging resolution, and the thickness of the transparent layer gradually increased, in most species with the transparent 1484:

from, these three types of cone cells. Cones are mostly concentrated in and near the fovea. Only a few are present at the sides of the retina. Objects are seen most sharply in focus when their images fall on the fovea, as when one looks at an object directly. Cone cells and rods are connected through

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of insects bear a simple lens, but their focal point usually lies behind the retina; consequently, those can not form a sharp image. Ocelli (pit-type eyes of arthropods) blur the image across the whole retina, and are consequently excellent at responding to rapid changes in light intensity across the

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Spherical aberration limits the resolution of a 7 mm pupil to about 3 arcminutes per line pair. At a pupil diameter of 3 mm, the spherical aberration is greatly reduced, resulting in an improved resolution of approximately 1.7 arcminutes per line pair. A resolution of 2 arcminutes per line

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a small ventral compound eye is physically completely separated from the much larger dorsal compound eye. Depth perception can be enhanced by having eyes which are enlarged in one direction; distorting the eye slightly allows the distance to the object to be estimated with a high degree of accuracy.

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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 produced by certain retinal cells. It is of rather similar composition to the cornea, but contains very few cells (mostly phagocytes which remove unwanted

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area which gives acute vision. In the acute zone, the eyes are flattened and the facets larger. The flattening allows more ommatidia to receive light from a spot and therefore higher resolution. The black spot that can be seen on the compound eyes of such insects, which always seems to look directly

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Through gradual change, the eye-spots of species living in well-lit environments depressed into a shallow "cup" shape. The ability to slightly discriminate directional brightness was achieved by using the angle at which the light hit certain cells to identify the source. The pit deepened over time,

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There are ten different eye layouts. Eye types can be categorised into "simple eyes", with one concave photoreceptive surface, and "compound eyes", which comprise a number of individual lenses laid out on a convex surface. "Simple" does not imply a reduced level of complexity or acuity. Indeed, any

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There are two types of opsin involved in vision; c-opsins, which are associated with ciliary-type photoreceptor cells, and r-opsins, associated with rhabdomeric photoreceptor cells. The eyes of vertebrates usually contain ciliary cells with c-opsins, and (bilaterian) invertebrates have rhabdomeric

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is triangular in horizontal section and is coated by a double layer, the ciliary epithelium. 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

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Apposition eyes work by gathering a number of images, one from each eye, and combining them in the brain, with each eye typically contributing a single point of information. The typical apposition eye has a lens focusing light from one direction on the rhabdom, while light from other directions is

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are composed of many simple facets which, depending on anatomical detail, may give either a single pixelated image or multiple images per eye. Each sensor has its own lens and photosensitive cell(s). Some eyes have up to 28,000 such sensors arranged hexagonally, which can give a full 360° field of

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1:posterior segment 2:ora serrata 3:ciliary muscle 4:ciliary zonules 5:Schlemm's canal 6:pupil 7:anterior chamber 8:cornea 9:iris 10:lens cortex 11:lens nucleus 12:ciliary process 13:conjunctiva 14:inferior oblique muscule 15:inferior rectus muscule 16:medial rectus muscle 17:retinal arteries and

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are deep water animals that feed on organisms above them. Their eyes are almost divided into two, with the upper region thought to be involved in detecting the silhouettes of potential prey—or predators—against the faint light of the sky above. Accordingly, deeper water hyperiids, where the light

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Of course, for most eye types, it is impossible to diverge from a spherical form, so only the density of optical receptors can be altered. In organisms with compound eyes, it is the number of ommatidia rather than ganglia that reflects the region of highest data acquisition. Optical superposition

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gene is considered a key factor in this. The majority of the advancements in early eyes are believed to have taken only a few million years to develop, since the first predator to gain true imaging would have touched off an "arms race" among all species that did not flee the photopic environment.

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The refracting superposition eye has a gap between the lens and the rhabdom, and no side wall. Each lens takes light at an angle to its axis and reflects it to the same angle on the other side. The result is an image at half the radius of the eye, which is where the tips of the rhabdoms are. This

869:, whose eyes include reflective optics for focusing of light. Each of the two eyes of a spookfish collects light from both above and below; the light coming from above is focused by a lens, while that coming from below, by a curved mirror composed of many layers of small reflective plates made of 688:

to form a lens, which may greatly reduce the blur radius encountered—hence increasing the resolution obtainable. The most basic form, seen in some gastropods and annelids, consists of a lens of one refractive index. A far sharper image can be obtained using materials with a high refractive index,

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Compound eyes fall into two groups: apposition eyes, which form multiple inverted images, and superposition eyes, which form a single erect image. Compound eyes are common in arthropods, annelids and some bivalved molluscs. Compound eyes in arthropods grow at their margins by the addition of new

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Most organisms with colour vision can detect ultraviolet light. This high energy light can be damaging to receptor cells. With a few exceptions (snakes, placental mammals), most organisms avoid these effects by having absorbent oil droplets around their cone cells. The alternative, developed by

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As mentioned above, a refractive cornea is only useful out of water. In water, there is little difference in refractive index between the vitreous fluid and the surrounding water. Hence creatures that have returned to the water—penguins and seals, for example—lose their highly curved cornea and

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into the brain on metamorphosis to the adult form. C-opsins are also found in some derived bilaterian-invertebrate eyes, such as the pallial eyes of the bivalve molluscs; however, the lateral eyes (which were presumably the ancestral type for this group, if eyes evolved once there) always use

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However, in the compound eye, the resolution is related to the size of individual ommatidia and the distance between neighbouring ommatidia. Physically these cannot be reduced in size to achieve the acuity seen with single lensed eyes as in mammals. Compound eyes have a much lower acuity than

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Acuity is higher among male organisms that mate in mid-air, as they need to be able to spot and assess potential mates against a very large backdrop. On the other hand, the eyes of organisms which operate in low light levels, such as around dawn and dusk or in deep water, tend to be larger to

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There are some exceptions from the types mentioned above. Some insects have a so-called single lens compound eye, a transitional type which is something between a superposition type of the multi-lens compound eye and the single lens eye found in animals with simple eyes. Then there is the

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The thin overgrowth of transparent cells over the eye's aperture, originally formed to prevent damage to the eyespot, allowed the segregated contents of the eye chamber to specialise into a transparent humour that optimised colour filtering, blocked harmful radiation, improved the eye's

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An alternative to a lens is to line the inside of the eye with "mirrors", and reflect the image to focus at a central point. The nature of these eyes means that if one were to peer into the pupil of an eye, one would see the same image that the organism would see, reflected back out.

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Eyes are generally adapted to the environment and life requirements of the organism which bears them. For instance, the distribution of photoreceptors tends to match the area in which the highest acuity is required, with horizon-scanning organisms, such as those that live on the

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in unicellular animals. In multicellular beings, multicellular eyespots evolved, physically similar to the receptor patches for taste and smell. These eyespots could only sense ambient brightness: they could distinguish light and dark, but not the direction of the light source.

1533:, which are an outgroup to the taxa mentioned above, express c-opsins—but r-opsins are yet to be found in this group. Incidentally, the melanin produced in the cnidaria is produced in the same fashion as that in vertebrates, suggesting the common descent of this pigment. 1060:. Because each eyelet is a simple eye, it produces an inverted image; those images are combined in the brain to form one unified image. Because the aperture of an eyelet is larger than the facets of a compound eye, this arrangement allows vision under low light levels. 1200:) of all animal eyes is now widely accepted as fact. This is based upon the shared genetic features of all eyes; that is, all modern eyes, varied as they are, have their origins in a proto-eye believed to have evolved some 650-600 million years ago, and the 798:), or with an aspheric shape. Flattening the lens has a disadvantage; the quality of vision is diminished away from the main line of focus. Thus, animals that have evolved with a wide field-of-view often have eyes that make use of an inhomogeneous lens. 765:

Multiple lenses are seen in some hunters such as eagles and jumping spiders, which have a refractive cornea: these have a negative lens, enlarging the observed image by up to 50% over the receptor cells, thus increasing their optical resolution.

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per line pair, or a 0.35 mm line pair, at 1 m). A rat can resolve only about 1 to 2 CPD. A horse has higher acuity through most of the visual field of its eyes than a human has, but does not match the high acuity of the human eye's central

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decreasing to the edges; this decreases the focal length and thus allows a sharp image to form on the retina. This also allows a larger aperture for a given sharpness of image, allowing more light to enter the lens; and a flatter lens, reducing

973:, the horseshoe crab, and there are suggestions that other chelicerates developed their simple eyes by reduction from a compound starting point. (Some caterpillars appear to have evolved compound eyes from simple eyes in the opposite fashion.) 1297:

An extension of this concept is that the eyes of predators typically have a zone of very acute vision at their centre, to assist in the identification of prey. In deep water organisms, it may not be the centre of the eye that is enlarged. The

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crystals formed the lenses of their eyes. They differ in this from most other arthropods, which have soft eyes. The number of lenses in such an eye varied widely; some trilobites had only one while others had thousands of lenses per eye.

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Müller, CHG; Rosenberg, J; Richter, S; Meyer-Rochow, VB (2003). "The compound eye of Scutigera coleoptrata (Linnaeus, 1758) (Chilopoda; Notostigmophora): an ultrastructural re-investigation that adds support to the Mandibulata concept".

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The pigment molecules used in the eye are various, but can be used to define the evolutionary distance between different groups, and can also be an aid in determining which are closely related—although problems of convergence do exist.

1489:. When rods and cones are stimulated by light, they connect through adjoining cells within the retina to send an electrical signal to the optic nerve fibres. The optic nerves send off impulses through these fibres to the brain. 1289:

plains, having a horizontal line of high-density ganglia, while tree-dwelling creatures which require good all-round vision tend to have a symmetrical distribution of ganglia, with acuity decreasing outwards from the centre.

564:, so are better suited to dark-dwelling creatures. Eyes also fall into two groups on the basis of their photoreceptor's cellular construction, with the photoreceptor cells either being ciliated (as in the vertebrates) or 1263:

The gap between tissue layers naturally formed a biconvex shape, an optimally ideal structure for a normal refractive index. Independently, a transparent layer and a nontransparent layer split forward from the lens: the

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It is not only the shape of the eye that may be affected by lifestyle. Eyes can be the most visible parts of organisms, and this can act as a pressure on organisms to have more transparent eyes at the cost of function.

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organisms that had lost these oil droplets in the course of evolution, is to make the lens impervious to UV light—this precludes the possibility of any UV light being detected, as it does not even reach the retina.

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evolved. Even if organisms are physically capable of discriminating different colours, this does not necessarily mean that they can perceive the different colours; only with behavioural tests can this be deduced.

1138:. The tube feet of sea urchins contain photoreceptor proteins, which together act as a compound eye; they lack screening pigments, but can detect the directionality of light by the shadow cast by its opaque body. 4114:

veins 18:optic disc 19:dura mater 20:central retinal artery 21:central retinal vein 22:optic nerve 23:vorticose vein 24:bulbar sheath 25:macula 26:fovea 27:sclera 28:choroid 29:superior rectus muscle 30:retina

943:). This can only be countered by increasing lens size and number. To see with a resolution comparable to our simple eyes, humans would require very large compound eyes, around 11 metres (36 ft) in radius. 730:

This eye creates an image that is sharp enough that motion of the eye can cause significant blurring. To minimise the effect of eye motion while the animal moves, most such eyes have stabilising eye muscles.

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The simplest eyes are pit eyes. They are eye-spots which may be set into a pit to reduce the angle of light that enters and affects the eye-spot, to allow the organism to deduce the angle of incoming light.

1508:, are used to shield the photoreceptor cells from light leaking in from the sides. The opsin protein group evolved long before the last common ancestor of animals, and has continued to diversify since. 2112:

Zhukov, ZH; Borisseko, SL; Zieger, MV; Vakoliuk, IA; Meyer-Rochow, VB (2006). "The eye of the freshwater prosobranch gastropod Viviparus viviparus: ultrastructure, electrophysiology and behaviour".

452:, have compound eyes of only a few facets, each with a retina capable of creating an image. With each eye producing a different image, a fused, high-resolution image is produced in the brain. 1224:, despite their distant common ancestry. Phenotypic convergence of the geometry of cephalopod and most vertebrate eyes creates the impression that the vertebrate eye evolved from an imaging 1322:

Eyes may be mounted on stalks to provide better all-round vision, by lifting them above an organism's carapace; this also allows them to track predators or prey without moving the head.

1163:), 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 727:

lenses. No extant aquatic organisms possess homogeneous lenses; presumably the evolutionary pressure for a heterogeneous lens is great enough for this stage to be quickly "outgrown".

1949: 364:. The last common ancestor of animals possessed the biochemical toolkit necessary for vision, and more advanced eyes have evolved in 96% of animal species in six of the ~35 main 1276:. This increased refractive power and again eased circulatory problems. Formation of a nontransparent ring allowed more blood vessels, more circulation, and larger eye sizes. 519:

larvae) can possess resolving powers of 4 degrees of arc, be polarization-sensitive, and capable of increasing its absolute sensitivity at night by a factor of 1,000 or more.

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against which the silhouettes must be compared is dimmer, have larger "upper-eyes", and may lose the lower portion of their eyes altogether. In the giant Antarctic isopod

832:, multiple rows of highly innervated and sensitive hairs which grow from the eyelid margins to protect the eye from fine particles and small irritants such as insects. 1919: 2430: 2714:

Horváth, Gábor; Clarkson, Euan N.K. (1997). "Survey of modern counterparts of schizochroal trilobite eyes: Structural and functional similarities and differences".

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have developed pits that function as eyes by sensing thermal infra-red radiation, in addition to their optical wavelength eyes like those of other vertebrates (see

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So-called under-focused lens eyes, found in gastropods and polychaete worms, have eyes that are intermediate between lens-less cup eyes and real camera eyes. Also

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cells but no lens or other means of projecting an image onto those cells. They can distinguish between light and dark but no more, enabling them to avoid direct

1457:) vision; they work well in dim light as they contain a pigment, rhodopsin (visual purple), which is sensitive at low light intensity, but saturates at higher ( 762:, has two lenses in each eye, arranged like those in a telescope. Such arrangements are rare and poorly understood, but represent an alternative construction. 3308:

Meyer-Rochow, Victor Benno (1982). "The divided eye of the isopod Glyptonotus antarcticus: effects of unilateral dark adaptation and temperature elevation".

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This eye type functions by refracting light, then using a parabolic mirror to focus the image; it combines features of superposition and apposition eyes.

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has three. The outer has a parabolic surface, countering the effects of spherical aberration while allowing a sharp image to be formed. Another copepod,

4095: 2263: 261:. Non-compound eyes have a single lens and focus light onto the retina to form a single image. This type of eye is common in mammals, including humans. 2194: 4100: 2050:

Kozmik, Z.; Ruzickova, J.; Jonasova, K.; Matsumoto, Y.; Vopalensky, P.; Kozmikova, I.; Strnad, H.; Kawamura, S.; Piatigorsky, J.; et al. (2008).

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eye type can be adapted for almost any behaviour or environment. The only limitations specific to eye types are that of resolution—the physics of

1228:, but this is not the case, as the reversed roles of their respective ciliary and rhabdomeric opsin classes and different lens crystallins show. 4096: 4101: 4081: 3887: 3957: 3866: 3600: 3494: 3419: 3370: 2930: 1891: 1845: 1818: 143: 139: 3462: 1937: 661: 4087: 1544: 3508: 3433: 3384: 1859: 2765: 1897: 862:

has up to 100 millimetre-scale reflector eyes fringing the edge of its shell. It detects moving objects as they pass successive lenses.

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Staaislav I. Tomarev; Rina D. Zinovieva (1988). "Squid major lens polypeptides are homologous to glutathione S-transferases subunits".

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There is no universal consensus on the precise total number of phyla Animalia; the stated figure varies slightly from author to author.

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hyaluronic acid, and also a wide array of proteins in micro amounts. Amazingly, with so little solid matter, it tautly holds the eye.

1052:, employs a series of simple eyes—eyes having one opening that provides light for an entire image-forming retina. Several of these 4092: 3242: 2790: 3648: 268:

Eyes enable several photo response functions that are independent of vision. In an organism that has more complex eyes, retinal

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Apposition eyes are the most common form of eyes and are presumably the ancestral form of compound eyes. They are found in all

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used this pigment in vision, and that remnants survive in the eyes. Likewise, c-opsins have been found to be expressed in the

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cellular debris in the visual field, as well as the hyalocytes of Balazs of the surface of the vitreous, which reprocess the

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The upper limit (finest detail) visible with the human eye is about 50 cycles per degree,... (Fifth Edition, 2007, Page 94)

599:. These are not considered eyes because they lack enough structure to be considered an organ, and do not produce an image. 420:, regulating the amount of light that enters the eye and reducing aberrations when there is enough light. The eyes of most 4083: 2449: 1758: 693:. Such a non-homogeneous lens is necessary for the focal length to drop from about 4 times the lens radius, to 2.5 radii. 602:

Every technological method of capturing an optical image that humans commonly use occurs in nature, with the exception of

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return to lens-based vision. An alternative solution, borne by some divers, is to have a very strongly focusing cornea.

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Eye strips images of all but bare essentials before sending visual information to the brain, UC Berkeley research shows

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Mayer, G. (2006). "Structure and development of onychophoran eyes: What is the ancestral visual organ in arthropods?".

1481: 4501: 4470: 4462: 4142: 3113: 1640: 779: 592: 4102: 4093: 4090: 4297: 4287: 669: 657: 4099: 4098: 392:(for low-light contrasts) in the retina detect and convert light into neural signals which are transmitted to the 4082: 3950: 1620: 1008:, the parabolic surfaces of the inside of each facet focus light from a reflector to a sensor array. Long-bodied 2498: 4105: 3979: 1590: 816:

are a unique characteristic of most mammalian eyes, both of which are evolutionary features to protect the eye.

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of some invertebrates. They may have been expressed in ciliary cells of larval eyes, which were subsequently

4659: 4292: 4248: 4106: 2525:"Optics and phylogeny: is there an insight? The evolution of superposition eyes in the Decapoda (Crustacea)" 1307: 338:. In other organisms, particularly prey animals, eyes are located to maximise the field of view, such as in 277: 51: 2190: 1056:

together form the strepsipteran compound eye, which is similar to the 'schizochroal' compound eyes of some

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Halder, G.; Callaerts, P.; Gehring, W.J. (1995). "Induction of ectopic eyes by targeted expression of the

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have fixed lens shapes, and focusing is achieved by telescoping the lens in a similar manner to that of a

285: 166: 150: 134: 660:). However, pit organs are fitted with receptors rather different from photoreceptors, namely a specific 4608: 4413: 4337: 4275: 4263: 4218: 1477: 1232: 401: 47: 4108: 3032:

Tomarev, S.I.; Callaerts, P.; Kos, L.; Zinovieva, R.; Halder, G.; Gehring, W.; Piatigorsky, J. (1997).

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the opening diminished in size, and the number of photoreceptor cells increased, forming an effective

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impose a limit on the possible resolution that can be obtained (assuming that they do not function as

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are alone in having reflecting superposition eyes, which also have a transparent gap but use corner

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The second type is named the superposition eye. The superposition eye is divided into three types:

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and non-compound eyes. Compound eyes are made up of multiple small visual units, and are common on

4088: 1134:, is covered with ommatidia, turning its whole skin into a compound eye. The same is true of many 852:

use such organs, but these are too small to produce usable images. Some larger organisms, such as

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An in-depth treatment of retinal function, open to all but geared most towards graduate students.

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across the cornea to prevent dehydration. These eyelids are also supplemented by the presence of

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This article is about the organ present in many organisms. For the human organ specifically, see

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radiation. The different forms of eye in, for example, vertebrates and molluscs are examples of

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Comparative Physiology and Evolution of Vision in Invertebrates- A: Invertebrate Photoreceptors

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system that collects light from the surrounding environment, regulates its intensity through a

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Land, M.F. (1989). "The eyes of hyperiid amphipods: relations of optical structure to depth".

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Wagner, H.J.; Douglas, R.H.; Frank, T.M.; Roberts, N.W. & Partridge, J.C. (Jan 27, 2009).

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Simple eyes are rather ubiquitous, and lens-bearing eyes have evolved at least seven times in

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The resolution of pit eyes can be greatly improved by incorporating a material with a higher

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Goldsmith, T.H. (1990). "Optimization, Constraint, and History in the Evolution of Eyes".

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Eyes with resolving power have come in ten fundamentally different forms, classified into

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Meyer-Rochow, V.B. (1974). "Structure and function of the larval eye of the sawfly larva

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For a human eye with excellent acuity, the maximum theoretical resolution is 50 CPD (1.2

1349:, or resolving power, is "the ability to distinguish fine detail" and is the property of 1208:

Eyes in various animals show adaptation to their requirements. For example, the eye of a

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A refractive cornea type eye of a human. The cornea is the clear domed part covering the

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A compound eye may consist of thousands of individual photoreceptor units or ommatidia (

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Fischer, Robert E.; Tadic-Galeb, Biljana; Plympton, Rick (2000). Steve Chapman (ed.).

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Halder, G.; Callaerts, P.; Gehring, W.J. (1995). "New perspectives on eye evolution".

2445: 4648: 4561: 4496: 4333: 3738:

Frentiu, Francesca D.; Adriana D. Briscoe (2008). "A butterfly eye's view of birds".

3592: 3068: 3033: 2959: 2125: 1988: 1923: 1630: 1401: 1385: 1346: 1193: 891: 795: 787: 697: 553: 496: 474: 470: 243: 208: 188: 32: 3835: 3767: 3717: 3333: 3294: 2415: 2318: 2176: 2019:"Exceptional Variation on a Common Theme: the Evolution of Crustacean Compound Eyes" 1461:) intensities. Rods are distributed throughout the retina but there are none at the 448:

vision. Compound eyes are very sensitive to motion. Some arthropods, including many

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Another version is a compound eye often referred to as "pseudofaceted", as seen in

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that was capable of dimly distinguishing shapes. However, the ancestors of modern

16:

Organ that detects light and converts it into electro-chemical impulses in neurons

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on each side of the head, organised in a way that resembles a true compound eye.

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Ullrich-Luter, E.M.; Dupont, S.; Arboleda, E.; Hausen, H.; Arnone, M.I. (2011).

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Fernald, Russell D. (2006). "Casting a Genetic Light on the Evolution of Eyes".

1605: 1486: 1217: 1186: 1081: 936: 791: 673: 565: 508: 397: 239: 2059:

Proceedings of the National Academy of Sciences of the United States of America

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groups, although they may have evolved more than once within this phylum. Some

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produced by the hot vents, allowing the creatures to avoid being boiled alive.

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have eyes with a spherical lens, cornea and retina, but the vision is blurry.

653: 631: 623: 619: 421: 369: 258: 3914: 3585: 3504: 3380: 3109:"Evolution of the vertebrate eye: opsins, photoreceptors, retina and eye cup" 2689: 1779: 159: 4403: 4398: 4317: 3971: 3429: 3010: 2876: 2160: 2079: 1855: 1595: 1443: 1414: 1410: 1368: 1213: 1197: 1118: 1113: 1085: 1068: 956: 724: 603: 573: 480: 444: 429: 385: 376:, the eye allows light to enter and project onto a light-sensitive layer of 311: 155: 92: 20: 3827: 3759: 3325: 3144: 3058: 2895: 2778: 2665: 2624: 2494: 2407: 2310: 2168: 2098: 2052:"Assembly of the cnidarian camera-type eye from vertebrate-like components" 703:

Heterogeneous eyes have evolved at least nine times: four or more times in

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Good fliers such as flies or honey bees, or prey-catching insects such as

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cells that do nothing but detect whether the surroundings are light or

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fields of many organisms, especially predators, involve large areas of

204: 3636: 523:, some of the simplest eyes, are found in animals such as some of the 4568: 4478: 4232: 4018: 3990: 3181: 1504:

are the pigments involved in photoreception. Other pigments, such as

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of light. Because the individual lenses are so small, the effects of

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to produce vision. Such eyes are typically spheroid, filled with the

381: 365: 355: 339: 323: 315: 254: 215: 3126: 2374:"A Novel Vertebrate Eye Using Both Refractive and Reflective Optics" 2231: 1449:

Rods cannot distinguish colours, but are responsible for low-light (

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very old, with various theories of phylogenesis. The common origin (

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Wilson, M. (1978). "The functional organisation of locust ocelli".

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The very earliest "eyes", called eye-spots, were simple patches of

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has the world's most complex colour vision system. It has detailed

42:"Eyeball", "Eyes", and "Ocular" redirect here. For other uses, see 4208: 1652: 1501: 1334: 1180: 1101: 1017: 914: 906: 895: 825: 804: 773: 665: 524: 504: 454: 433: 417: 393: 343: 306: 295: 235: 231: 200: 172: 122: 28: 3924: 2640:"Dimensional limits for arthropod eyes with superposition optics" 4122: 1201: 747:

Some marine organisms bear more than one lens; for instance the

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prevents them from achieving a resolution better than 1°. Also,

425: 3939: 2754:"Variations in the optical properties of the compound eyes of 1920:"What animal has a more sophisticated eye, Octopus or Insect?" 1562:

Another view of the eye and the structures of the eye labelled

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In contrast to compound eyes, simple eyes have a single lens.

465:) are considered the most complex in the whole animal kingdom. 3411:

Casarett and Doull's Toxicology: The Basic Science of Poisons

1516:

cells of vertebrates express r-opsins, suggesting that their

1272:. Separation of the forward layer again formed a humour, the 4076: 3925:

Webvision. The organisation of the retina and visual system.

3107:

Trevor D. Lamb; Shaun P. Collin; Edward N. Pugh Jr. (2007).

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Scourfield, a protochordate from the Silurian of Scotland".

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Bruce, Vicki; Green, Patrick R.; Georgeson, Mark A. (1996).

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Land, M.F.; Fernald, R.D. (1992). "The evolution of eyes".

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Adaptations for nocturnal vision in insect apposition eyes

2852:"Unique system of photoreceptors in sea urchin tube feet" 24: 1485:

intermediate cells in the retina to nerve fibres of the

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A unique feature of most mammal eyes is the presence of

515:) which usually provides only a rough image, but (as in 2216:

Nilsson, Dan-E. (1989). "Vision optics and evolution".

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The retina contains two major types of light-sensitive

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Autrum, H (1979). "Introduction". In H. Autrum (ed.).

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Another kind of compound eye, found in males of Order

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also have apposition eyes. They are also possessed by

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An image of a house fly compound eye surface by using

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Speiser, D.I.; Eernisse, D.J.; Johnsen, S.N. (2011).

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Visual Perception: Physiology, Psychology and Ecology

1117:. This type of eye consists of a cluster of numerous 238:

through neural pathways that connect the eye via the

4586: 4547: 4461: 4452: 4386: 4306: 4241: 4231: 4194: 4161: 4133: 4121: 4017: 3989: 3978: 3234: 3232: 1315:increase the amount of light that can be captured. 149: 133: 121: 116: 104: 99: 61: 3888:"Inside the Eye: Nature's Most Exquisite Creation" 3584: 2193:. National Institute of General Medical Sciences. 1802: 203:and converts it into electro-chemical impulses in 3591:. Cambridge: Cambridge University Press. p. 3240:The Evolution of Eyes: Where Do Lenses Come From? 668:. The main difference is that photoreceptors are 495:have one pair of large simple eyes with a narrow 416:. Muscles around the iris change the size of the 3915:Anatomy of the eye – flash animated interactive. 3673: 3671: 3669: 3667: 3665: 4080: 2856:Proceedings of the National Academy of Sciences 2277:"A Chiton Uses Aragonite Lenses to Form Images" 2264:Perceiving in Depth, Volume 1: Basic Mechanisms 483:, now extinct, had unique compound eyes. Clear 3787: 3785: 3783: 3781: 3779: 3777: 3733: 3731: 3729: 3727: 2429:Völkel, R; Eisner, M; Weible, KJ (June 2003). 1512:cells in the eye with r-opsins. However, some 3951: 2332: 2330: 2328: 1809:. New York: Oxford University Press. p. 1151:constitutes the cells of the dilator muscle. 568:. These two groups are not monophyletic; the 8: 2012: 2010: 2008: 2006: 1380:pair, equivalent to a 1 arcminute gap in an 499:, augmented by an array of smaller eyes for 3583:Barlow, Horace Basil; Mollon, J.D. (1982). 3268: 3266: 3264: 3262: 3260: 3258: 1751: 1749: 1747: 1745: 1743: 1741: 1739: 1737: 1735: 1733: 1731: 1729: 1727: 1725: 1723: 1721: 1719: 1717: 1715: 1713: 353:The first proto-eyes evolved among animals 4458: 4238: 4130: 3986: 3958: 3944: 3936: 3531:"The size of ommatidia in apposition eyes" 3310:Proceedings of the Royal Society of London 1711: 1709: 1707: 1705: 1703: 1701: 1699: 1697: 1695: 1693: 1088:which one observes "head-on" (along their 280:to effect circadian adjustment and to the 214:In higher organisms, the eye is a complex 85: 67: 3817: 3134: 3067: 3057: 2885: 2875: 2655: 2540: 2397: 2300: 2088: 2078: 2034: 3853:Ali, Mohamed Ather; Klyne, M.A. (1985). 3570: 3346: 3214:. Library.thinkquest.org. Archived from 2908: 2752:Jochen Zeil; Maha M. Al-Mutairi (1996). 911:Anatomy of the compound eye of an insect 207:(neurones). It is part of an organism's 3102: 3100: 2747: 2745: 1805:Blindspots: The Many Ways We Cannot See 1689: 1664: 1540: 856:, also use reflector eyes. The scallop 572:also possess ciliated cells, and some 557: 539:, compound eyes are adapted to see the 2688:(PhD). Lund University. Archived from 1212:has much greater visual acuity than a 1096:, while those to one side reflect it. 865:There is at least one vertebrate, the 170: 58: 3794:"Eye Evolution: the Blurry Beginning" 1922:. BioMedia Associates. Archived from 560:can achieve greater sensitivity than 226:it through an adjustable assembly of 7: 3792:Nilsson, E.; Arendt, D. (Dec 2008). 2679:Greiner, Birgit (16 December 2005). 1918:Kirk, Molly; Denning, David (2001). 662:transient receptor potential channel 322:Complex eyes distinguish shapes and 3275:Journal of Comparative Physiology A 2766:The Journal of Experimental Biology 2597:Arthropod Structure and Development 2017:Cronin, T.W.; Porter, M.L. (2008). 1772:10.1146/annurev.ne.15.030192.000245 2197:from the original on 13 March 2020 1952:from the original on 9 August 2010 1550:The structures of the eye labelled 459:The eyes of a mantis shrimp (here 14: 3094:. Oxford: Oxford University Press 3034:"Squid Pax-6 and eye development" 2638:Meyer-Rochow, VB; Gal, J (2004). 2339:Journal of Comparative Physiology 2023:Evolution: Education and Outreach 1840:. Belmont: Wadsworth Publishing. 1280:Relationship to life requirements 977:absorbed by the dark wall of the 2126:10.1111/j.1463-6395.2006.00216.x 1886:. Psychology Press. p. 20. 1555: 1543: 1417:, from which the more sensitive 3681:The Quarterly Review of Biology 3651:from the original on 2012-11-20 3553:from the original on 2016-08-31 3511:from the original on 2023-01-17 3465:from the original on 2015-05-04 3436:from the original on 2023-01-17 3387:from the original on 2023-01-17 2796:from the original on 2009-02-25 1900:from the original on 2023-01-17 1862:from the original on 2023-01-17 1216:, and in some cases can detect 31:. For region of a cyclone, see 2431:"Miniaturized imaging systems" 2191:"Circadian Rhythms Fact Sheet" 1574:Accommodation (vertebrate eye) 824:which wipe the eye and spread 591:, which is sufficient for the 246:and other areas of the brain. 1: 3625:Brain, Behavior and Evolution 3459:hyperphysics.phy-astr.gsu.edu 3455:"The Retina of the Human Eye" 3362:The Image Processing Handbook 2487:10.1146/annurev.ento.42.1.147 2446:10.1016/S0167-9317(03)00102-3 1759:Annual Review of Neuroscience 1080:at the observer, is called a 844:Many small organisms such as 535:. In organisms dwelling near 511:, have a type of simple eye ( 270:photosensitive ganglion cells 4604:Optical coherence tomography 4358:Photosensitive ganglion cell 3616:Fernald, Russell D. (1997). 3489:. McGraw-Hill Professional. 3414:. McGraw-Hill Professional. 3408:Klaassen, Curtis D. (2001). 3238:Fernald, Russell D. (2001). 2960:10.1016/0959-437X(95)80029-8 2657:10.1016/j.visres.2004.04.009 2556:Ritchie, Alexander (1985). " 1989:10.1016/0022-1910(74)90087-0 1977:Journal of Insect Physiology 1946:National Wildlife Federation 1409:The most sensitive pigment, 1071:, have specialised zones of 902:scanning electron microscope 4354:Giant retina ganglion cells 4143:Capillary lamina of choroid 3114:Nature Reviews Neuroscience 2475:Annual Review of Entomology 2438:Microelectronic Engineering 1938:"Who You Callin' "Shrimp"?" 1641:Simple eye in invertebrates 780:anterior chamber of the eye 670:G-protein coupled receptors 4681: 4298:Retinal pigment epithelium 4288:External limiting membrane 3090:Conway-Morris, S. (1998). 3038:Proc. Natl. Acad. Sci. USA 2468:"Visual Acuity in Insects" 1942:National Wildlife Magazine 1801:Breitmeyer, Bruno (2010). 1472:Cones are responsible for 1399: 1353:. It is often measured in 1174: 1084:. This occurs because the 889: 883: 658:infrared sensing in snakes 41: 18: 4074: 3819:10.1016/j.cub.2008.10.025 3312:. B 215 (1201): 433–450. 2829:10.1007/s00435-003-0085-0 2736:10.1080/08912969709386565 2617:10.1016/j.asd.2006.06.003 2582:10.1080/03115518508618961 2542:10.1163/18759866-06704001 2399:10.1016/j.cub.2008.11.061 2302:10.1016/j.cub.2011.03.033 2036:10.1007/s12052-008-0085-0 1621:Lens (vertebrate anatomy) 1360:(CPD), which measures an 1106:Dioptromysis paucispinosa 443:The compound eyes of the 165: 84: 66: 3092:The Crucible of Creation 2529:Contributions to Zoology 1591:Emission theory (vision) 1384:, corresponds to 20/20 ( 919:Arthropods such as this 646:Pit eyes, also known as 462:Odontodactylus scyllarus 274:retinohypothalamic tract 199:information. It detects 44:Eyeball (disambiguation) 4293:Layer of rods and cones 4249:Inner limiting membrane 3618:"The Evolution of Eyes" 3252:64: "The Eye in Focus". 3011:10.1126/science.7892602 2877:10.1073/pnas.1018495108 2161:10.1126/science.1127889 2080:10.1073/pnas.0800388105 1036:Parabolic superposition 999:parabolic superposition 358: million years ago 272:send signals along the 52:Ocular (disambiguation) 4115: 3359:Russ, John C. (2006). 3326:10.1098/rspb.1982.0052 3059:10.1073/pnas.94.6.2421 2948:Curr. Opin. Genet. Dev 2779:10.1242/jeb.199.7.1569 2523:Gaten, Edward (1998). 2466:Land, Michael (1997). 2440:. 67–68 (1): 461–472. 1834:Nairne, James (2005). 1343: 1189: 924: 912: 904: 817: 783: 664:(TRP channels) called 466: 360:about the time of the 319: 304: 286:pupillary light reflex 278:suprachiasmatic nuclei 167:Anatomical terminology 35:. For other uses, see 23:. For the letter, see 4609:Eye care professional 4414:Foveal avascular zone 4276:Outer plexiform layer 4264:Inner plexiform layer 4219:Iris sphincter muscle 4112: 3855:Vision in Vertebrates 3529:Barlow, H.B. (1952). 3486:Optical System Design 2558:Ainiktozoon loganense 1438:used for vision: the 1338: 1233:photoreceptor protein 1184: 918: 910: 899: 890:Further information: 808: 777: 458: 408:, possess a focusing 388:(for colour) and the 310: 299: 48:Eyes (disambiguation) 27:. For the pupil, see 4629:Physiological Optics 4599:Ocular immune system 4338:Retina ganglion cell 3910:Evolution of the eye 3898:on January 14, 2016. 3746:(11–12): 1151–1162. 3571:Ali & Klyne 1985 3547:10.1242/jeb.29.4.667 3347:Ali & Klyne 1985 2909:Ali & Klyne 1985 2756:Uca lactea annulipes 1926:on 26 February 2017. 1626:Nictitating membrane 1177:Evolution of the eye 788:eyes of most mammals 691:spherical aberration 583:Some organisms have 406:vitreous humour 37:Eye (disambiguation) 4453:Anatomical regions 4314:Photoreceptor cells 4281:Outer nuclear layer 4269:Inner nuclear layer 4259:Ganglion cell layer 4214:Iris dilator muscle 4009:Trabecular meshwork 3892:National Geographic 3886:(14 January 2016). 3810:2008CBio...18R1096N 3804:(23): R1096–R1098. 3318:1982RSPSB.215..433M 3174:1988Natur.336...86T 3050:1997PNAS...94.2421T 3003:1995Sci...267.1788H 2997:(5205): 1788–1792. 2868:2011PNAS..108.8367U 2728:1997HBio...12..229H 2609:2006ArtSD..35..231M 2574:1985Alch....9..117R 2504:on 23 November 2004 2390:2009CBio...19..108W 2293:2011CBio...21..665S 2153:2006Sci...313.1914F 2147:(5795): 1914–1918. 2071:2008PNAS..105.8989K 1436:photoreceptor cells 1032:instead of lenses. 1010:decapod crustaceans 923:have compound eyes. 4116: 3752:10.1002/bies.20828 3287:10.1007/BF00616747 3245:2006-03-19 at the 2716:Historical Biology 2695:on 9 February 2013 2351:10.1007/BF00661380 1948:. 1 October 2010. 1362:angular resolution 1344: 1222:parallel evolution 1192:Photoreception is 1190: 1165:mucopolysaccharide 985:Superposition eyes 925: 913: 905: 818: 784: 719:, and once in the 680:Spherical lens eye 558:superposition eyes 467: 362:Cambrian explosion 320: 305: 4642: 4641: 4634:Visual perception 4582: 4581: 4549:Posterior segment 4517:Posterior chamber 4448: 4447: 4350:Bistratified cell 4254:Nerve fiber layer 4227: 4226: 4171:Ciliary processes 4072: 4071: 3868:978-0-306-42065-8 3637:10.1159/000113339 3602:978-0-521-24474-9 3496:978-0-07-134916-1 3421:978-0-07-134721-1 3372:978-0-8493-7254-4 2932:978-3-540-08837-0 2862:(20): 8367–8372. 2650:(19): 2213–2223. 2065:(26): 8989–8993. 1893:978-0-86377-450-8 1847:978-0-495-03150-5 1820:978-0-19-539426-9 1537:Additional images 1396:Colour perception 1392:vertebrate eyes. 1185:Evolution of the 1127:Ophiocoma wendtii 1075:organised into a 770:Refractive cornea 614:Non-compound eyes 597:circadian rhythms 501:peripheral vision 181: 180: 176: 4672: 4619:Refractive error 4557:Vitreous chamber 4502:Anterior chamber 4463:Anterior segment 4459: 4239: 4148:Bruch's membrane 4131: 4123:Uvea / vascular 4079: 3999:Episcleral layer 3987: 3960: 3953: 3946: 3937: 3899: 3894:. Archived from 3872: 3840: 3839: 3821: 3789: 3772: 3771: 3735: 3722: 3721: 3675: 3660: 3659: 3657: 3656: 3622: 3613: 3607: 3606: 3590: 3580: 3574: 3568: 3562: 3561: 3559: 3558: 3526: 3520: 3519: 3517: 3516: 3480: 3474: 3473: 3471: 3470: 3451: 3445: 3444: 3442: 3441: 3405: 3399: 3398: 3393: 3392: 3356: 3350: 3344: 3338: 3337: 3305: 3299: 3298: 3270: 3253: 3236: 3227: 3226: 3224: 3223: 3212:"Eye-Evolution?" 3208: 3202: 3201: 3182:10.1038/336086a0 3155: 3149: 3148: 3138: 3104: 3095: 3088: 3082: 3081: 3071: 3061: 3044:(6): 2421–2426. 3029: 3023: 3022: 2978: 2972: 2971: 2943: 2937: 2936: 2918: 2912: 2906: 2900: 2899: 2889: 2879: 2847: 2841: 2840: 2811: 2805: 2804: 2802: 2801: 2795: 2773:(7): 1569–1577. 2762: 2749: 2740: 2739: 2722:(3–4): 229–263. 2711: 2705: 2704: 2702: 2700: 2694: 2687: 2676: 2670: 2669: 2659: 2635: 2629: 2628: 2592: 2586: 2585: 2553: 2547: 2546: 2544: 2520: 2514: 2513: 2511: 2509: 2503: 2497:. Archived from 2472: 2463: 2457: 2456: 2454: 2448:. Archived from 2435: 2426: 2420: 2419: 2401: 2369: 2363: 2362: 2334: 2323: 2322: 2304: 2272: 2266: 2261: 2255: 2250: 2244: 2243: 2213: 2207: 2206: 2204: 2202: 2187: 2181: 2180: 2136: 2130: 2129: 2109: 2103: 2102: 2092: 2082: 2056: 2055:(Free full text) 2047: 2041: 2040: 2038: 2014: 2001: 2000: 1983:(8): 1565–1591. 1968: 1962: 1961: 1959: 1957: 1934: 1928: 1927: 1915: 1909: 1908: 1906: 1905: 1877: 1871: 1870: 1868: 1867: 1831: 1825: 1824: 1808: 1798: 1792: 1791: 1753: 1672: 1669: 1580:Adaptation (eye) 1559: 1547: 1465:and none at the 1254:refractive index 1194:phylogenetically 686:refractive index 359: 348:monocular vision 336:depth perception 332:binocular vision 173:edit on Wikidata 89: 71: 59: 4680: 4679: 4675: 4674: 4673: 4671: 4670: 4669: 4645: 4644: 4643: 4638: 4578: 4543: 4532:Capsule of lens 4487:Lacrimal system 4454: 4444: 4404:Parafoveal area 4399:Perifoveal area 4382: 4326:Horizontal cell 4302: 4223: 4190: 4157: 4153:Sattler's layer 4124: 4117: 4111: 4068: 4013: 4004:Schlemm's canal 3982: 3974: 3966:Anatomy of the 3964: 3906: 3882: 3879: 3877:Further reading 3869: 3852: 3849: 3844: 3843: 3798:Current Biology 3791: 3790: 3775: 3737: 3736: 3725: 3677: 3676: 3663: 3654: 3652: 3620: 3615: 3614: 3610: 3603: 3582: 3581: 3577: 3569: 3565: 3556: 3554: 3528: 3527: 3523: 3514: 3512: 3497: 3482: 3481: 3477: 3468: 3466: 3453: 3452: 3448: 3439: 3437: 3422: 3407: 3406: 3402: 3390: 3388: 3373: 3358: 3357: 3353: 3345: 3341: 3307: 3306: 3302: 3272: 3271: 3256: 3247:Wayback Machine 3237: 3230: 3221: 3219: 3210: 3209: 3205: 3168:(6194): 86–88. 3157: 3156: 3152: 3127:10.1038/nrn2283 3121:(12): 960–976. 3106: 3105: 3098: 3089: 3085: 3031: 3030: 3026: 2980: 2979: 2975: 2945: 2944: 2940: 2933: 2920: 2919: 2915: 2907: 2903: 2849: 2848: 2844: 2813: 2812: 2808: 2799: 2797: 2793: 2760: 2751: 2750: 2743: 2713: 2712: 2708: 2698: 2696: 2692: 2685: 2678: 2677: 2673: 2644:Vision Research 2637: 2636: 2632: 2594: 2593: 2589: 2555: 2554: 2550: 2522: 2521: 2517: 2507: 2505: 2501: 2470: 2465: 2464: 2460: 2452: 2433: 2428: 2427: 2423: 2378:Current Biology 2371: 2370: 2366: 2336: 2335: 2326: 2281:Current Biology 2274: 2273: 2269: 2262: 2258: 2251: 2247: 2232:10.2307/1311112 2215: 2214: 2210: 2200: 2198: 2189: 2188: 2184: 2138: 2137: 2133: 2111: 2110: 2106: 2054: 2049: 2048: 2044: 2016: 2015: 2004: 1970: 1969: 1965: 1955: 1953: 1936: 1935: 1931: 1917: 1916: 1912: 1903: 1901: 1894: 1879: 1878: 1874: 1865: 1863: 1848: 1833: 1832: 1828: 1821: 1800: 1799: 1795: 1755: 1754: 1691: 1686: 1681: 1676: 1675: 1670: 1666: 1661: 1647:Tapetum lucidum 1636:Orbit (anatomy) 1601:Eye development 1586:Capsule of lens 1570: 1563: 1560: 1551: 1548: 1539: 1495: 1455:black-and-white 1432: 1404: 1398: 1341:red-tailed hawk 1333: 1328: 1282: 1179: 1173: 1161:hyaluronic acid 1144: 1046: 1038: 987: 953: 951:Apposition eyes 921:blue bottle fly 894: 888: 882: 848:, copepods and 838: 772: 745: 743:Multiple lenses 682: 644: 616: 562:apposition eyes 549: 541:infra-red light 493:Jumping spiders 477:colour vision. 412:, and often an 354: 314:, a refractive 294: 284:to control the 191:that allows an 177: 142: 95: 80: 78:Antarctic krill 55: 40: 17: 12: 11: 5: 4678: 4676: 4668: 4667: 4662: 4660:Sensory organs 4657: 4647: 4646: 4640: 4639: 4637: 4636: 4631: 4626: 4621: 4616: 4611: 4606: 4601: 4596: 4590: 4588: 4584: 4583: 4580: 4579: 4577: 4576: 4571: 4566: 4565: 4564: 4553: 4551: 4545: 4544: 4542: 4541: 4540: 4539: 4537:Zonule of Zinn 4534: 4524: 4519: 4514: 4509: 4507:Aqueous humour 4504: 4499: 4494: 4467: 4465: 4456: 4450: 4449: 4446: 4445: 4443: 4442: 4437: 4436: 4435: 4425: 4424: 4423: 4422: 4421: 4416: 4406: 4401: 4390: 4388: 4384: 4383: 4381: 4380: 4310: 4308: 4304: 4303: 4301: 4300: 4295: 4290: 4284: 4283: 4278: 4272: 4271: 4266: 4261: 4256: 4251: 4245: 4243: 4236: 4229: 4228: 4225: 4224: 4222: 4221: 4216: 4211: 4206: 4200: 4198: 4192: 4191: 4189: 4188: 4183: 4178: 4176:Ciliary muscle 4173: 4167: 4165: 4159: 4158: 4156: 4155: 4150: 4145: 4139: 4137: 4128: 4119: 4118: 4075: 4073: 4070: 4069: 4067: 4066: 4065: 4064: 4059: 4054: 4049: 4044: 4039: 4029: 4023: 4021: 4015: 4014: 4012: 4011: 4006: 4001: 3995: 3993: 3984: 3976: 3975: 3965: 3963: 3962: 3955: 3948: 3940: 3934: 3933: 3928: 3922: 3912: 3905: 3904:External links 3902: 3901: 3900: 3878: 3875: 3874: 3873: 3867: 3848: 3845: 3842: 3841: 3773: 3723: 3694:10.1086/416840 3688:(3): 281–322. 3661: 3631:(4): 253–259. 3608: 3601: 3575: 3563: 3541:(4): 667–674. 3521: 3495: 3475: 3446: 3420: 3400: 3371: 3351: 3339: 3300: 3281:(6): 751–762. 3254: 3250:Karger Gazette 3228: 3203: 3150: 3096: 3083: 3024: 2973: 2954:(5): 602–609. 2938: 2931: 2913: 2901: 2842: 2823:(4): 191–209. 2806: 2741: 2706: 2671: 2630: 2603:(4): 231–245. 2587: 2548: 2535:(4): 223–236. 2515: 2458: 2455:on 2008-10-01. 2421: 2384:(2): 108–114. 2364: 2345:(4): 297–316. 2324: 2287:(8): 665–670. 2267: 2256: 2245: 2226:(5): 298–307. 2208: 2182: 2131: 2114:Acta Zoologica 2104: 2042: 2029:(4): 463–475. 2002: 1963: 1929: 1910: 1892: 1872: 1846: 1826: 1819: 1793: 1688: 1687: 1685: 1682: 1680: 1677: 1674: 1673: 1663: 1662: 1660: 1657: 1656: 1655: 1650: 1643: 1638: 1633: 1628: 1623: 1618: 1613: 1608: 1603: 1598: 1593: 1588: 1583: 1582:(night vision) 1577: 1569: 1566: 1565: 1564: 1561: 1554: 1552: 1549: 1542: 1538: 1535: 1494: 1491: 1453:) monochrome ( 1431: 1430:Rods and cones 1428: 1400:Main article: 1397: 1394: 1332: 1329: 1327: 1324: 1281: 1278: 1274:aqueous humour 1242:pinhole camera 1226:cephalopod eye 1175:Main article: 1172: 1169: 1143: 1140: 1094:incident light 1065:praying mantis 1045: 1042: 1037: 1034: 1001: 1000: 997: 996:reflecting and 994: 986: 983: 952: 949: 884:Main article: 881: 878: 837: 836:Reflector eyes 834: 771: 768: 744: 741: 715:, once in the 711:, once in the 707:, once in the 681: 678: 643: 640: 615: 612: 608:Fresnel lenses 585:photosensitive 580:possess both. 548: 545: 537:deep-sea vents 529:photosensitive 503:. Some insect 302:European bison 293: 290: 282:pretectal area 179: 178: 169: 163: 162: 153: 147: 146: 137: 131: 130: 125: 119: 118: 114: 113: 108: 102: 101: 97: 96: 90: 82: 81: 72: 64: 63: 15: 13: 10: 9: 6: 4: 3: 2: 4677: 4666: 4665:Visual system 4663: 4661: 4658: 4656: 4653: 4652: 4650: 4635: 4632: 4630: 4627: 4625: 4624:Accommodation 4622: 4620: 4617: 4615: 4612: 4610: 4607: 4605: 4602: 4600: 4597: 4595: 4592: 4591: 4589: 4585: 4575: 4572: 4570: 4567: 4563: 4562:Vitreous body 4560: 4559: 4558: 4555: 4554: 4552: 4550: 4546: 4538: 4535: 4533: 4530: 4529: 4528: 4525: 4523: 4520: 4518: 4515: 4513: 4510: 4508: 4505: 4503: 4500: 4498: 4497:Fibrous tunic 4495: 4492: 4488: 4484: 4480: 4476: 4472: 4469: 4468: 4466: 4464: 4460: 4457: 4451: 4441: 4438: 4434: 4431: 4430: 4429: 4426: 4420: 4417: 4415: 4412: 4411: 4410: 4407: 4405: 4402: 4400: 4397: 4396: 4395: 4392: 4391: 4389: 4385: 4379: 4375: 4371: 4367: 4363: 4359: 4355: 4351: 4347: 4343: 4339: 4335: 4334:Amacrine cell 4331: 4327: 4323: 4319: 4315: 4312: 4311: 4309: 4305: 4299: 4296: 4294: 4291: 4289: 4286: 4285: 4282: 4279: 4277: 4274: 4273: 4270: 4267: 4265: 4262: 4260: 4257: 4255: 4252: 4250: 4247: 4246: 4244: 4240: 4237: 4234: 4230: 4220: 4217: 4215: 4212: 4210: 4207: 4205: 4202: 4201: 4199: 4197: 4193: 4187: 4184: 4182: 4179: 4177: 4174: 4172: 4169: 4168: 4166: 4164: 4160: 4154: 4151: 4149: 4146: 4144: 4141: 4140: 4138: 4136: 4132: 4129: 4126: 4120: 4063: 4060: 4058: 4055: 4053: 4050: 4048: 4045: 4043: 4040: 4038: 4035: 4034: 4033: 4030: 4028: 4025: 4024: 4022: 4020: 4016: 4010: 4007: 4005: 4002: 4000: 3997: 3996: 3994: 3992: 3988: 3985: 3981: 3980:Fibrous tunic 3977: 3973: 3969: 3961: 3956: 3954: 3949: 3947: 3942: 3941: 3938: 3932: 3929: 3926: 3923: 3920: 3916: 3913: 3911: 3908: 3907: 3903: 3897: 3893: 3889: 3885: 3881: 3880: 3876: 3870: 3864: 3860: 3856: 3851: 3850: 3846: 3837: 3833: 3829: 3825: 3820: 3815: 3811: 3807: 3803: 3799: 3795: 3788: 3786: 3784: 3782: 3780: 3778: 3774: 3769: 3765: 3761: 3757: 3753: 3749: 3745: 3741: 3734: 3732: 3730: 3728: 3724: 3719: 3715: 3711: 3707: 3703: 3699: 3695: 3691: 3687: 3683: 3682: 3674: 3672: 3670: 3668: 3666: 3662: 3650: 3646: 3642: 3638: 3634: 3630: 3626: 3619: 3612: 3609: 3604: 3598: 3594: 3589: 3588: 3579: 3576: 3573:, p. 161 3572: 3567: 3564: 3552: 3548: 3544: 3540: 3536: 3532: 3525: 3522: 3510: 3506: 3502: 3498: 3492: 3488: 3487: 3479: 3476: 3464: 3460: 3456: 3450: 3447: 3435: 3431: 3427: 3423: 3417: 3413: 3412: 3404: 3401: 3397: 3386: 3382: 3378: 3374: 3368: 3365:. CRC Press. 3364: 3363: 3355: 3352: 3348: 3343: 3340: 3335: 3331: 3327: 3323: 3319: 3315: 3311: 3304: 3301: 3296: 3292: 3288: 3284: 3280: 3276: 3269: 3267: 3265: 3263: 3261: 3259: 3255: 3251: 3248: 3244: 3241: 3235: 3233: 3229: 3218:on 2012-09-15 3217: 3213: 3207: 3204: 3199: 3195: 3191: 3187: 3183: 3179: 3175: 3171: 3167: 3163: 3162: 3154: 3151: 3146: 3142: 3137: 3132: 3128: 3124: 3120: 3116: 3115: 3110: 3103: 3101: 3097: 3093: 3087: 3084: 3079: 3075: 3070: 3065: 3060: 3055: 3051: 3047: 3043: 3039: 3035: 3028: 3025: 3020: 3016: 3012: 3008: 3004: 3000: 2996: 2992: 2988: 2984: 2977: 2974: 2969: 2965: 2961: 2957: 2953: 2949: 2942: 2939: 2934: 2928: 2924: 2917: 2914: 2910: 2905: 2902: 2897: 2893: 2888: 2883: 2878: 2873: 2869: 2865: 2861: 2857: 2853: 2846: 2843: 2838: 2834: 2830: 2826: 2822: 2818: 2817:Zoomorphology 2810: 2807: 2792: 2788: 2784: 2780: 2776: 2772: 2768: 2767: 2759: 2757: 2748: 2746: 2742: 2737: 2733: 2729: 2725: 2721: 2717: 2710: 2707: 2691: 2684: 2683: 2675: 2672: 2667: 2663: 2658: 2653: 2649: 2645: 2641: 2634: 2631: 2626: 2622: 2618: 2614: 2610: 2606: 2602: 2598: 2591: 2588: 2583: 2579: 2575: 2571: 2567: 2563: 2559: 2552: 2549: 2543: 2538: 2534: 2530: 2526: 2519: 2516: 2500: 2496: 2492: 2488: 2484: 2480: 2476: 2469: 2462: 2459: 2451: 2447: 2443: 2439: 2432: 2425: 2422: 2417: 2413: 2409: 2405: 2400: 2395: 2391: 2387: 2383: 2379: 2375: 2368: 2365: 2360: 2356: 2352: 2348: 2344: 2340: 2333: 2331: 2329: 2325: 2320: 2316: 2312: 2308: 2303: 2298: 2294: 2290: 2286: 2282: 2278: 2271: 2268: 2265: 2260: 2257: 2254: 2249: 2246: 2241: 2237: 2233: 2229: 2225: 2221: 2220: 2212: 2209: 2196: 2192: 2186: 2183: 2178: 2174: 2170: 2166: 2162: 2158: 2154: 2150: 2146: 2142: 2135: 2132: 2127: 2123: 2119: 2115: 2108: 2105: 2100: 2096: 2091: 2086: 2081: 2076: 2072: 2068: 2064: 2060: 2053: 2046: 2043: 2037: 2032: 2028: 2024: 2020: 2013: 2011: 2009: 2007: 2003: 1998: 1994: 1990: 1986: 1982: 1978: 1974: 1967: 1964: 1951: 1947: 1943: 1939: 1933: 1930: 1925: 1921: 1914: 1911: 1899: 1895: 1889: 1885: 1884: 1876: 1873: 1861: 1857: 1853: 1849: 1843: 1839: 1838: 1830: 1827: 1822: 1816: 1812: 1807: 1806: 1797: 1794: 1789: 1785: 1781: 1777: 1773: 1769: 1765: 1761: 1760: 1752: 1750: 1748: 1746: 1744: 1742: 1740: 1738: 1736: 1734: 1732: 1730: 1728: 1726: 1724: 1722: 1720: 1718: 1716: 1714: 1712: 1710: 1708: 1706: 1704: 1702: 1700: 1698: 1696: 1694: 1690: 1683: 1678: 1668: 1665: 1658: 1654: 1651: 1649: 1648: 1644: 1642: 1639: 1637: 1634: 1632: 1631:Ophthalmology 1629: 1627: 1624: 1622: 1619: 1617: 1614: 1612: 1609: 1607: 1604: 1602: 1599: 1597: 1594: 1592: 1589: 1587: 1584: 1581: 1578: 1575: 1572: 1571: 1567: 1558: 1553: 1546: 1541: 1536: 1534: 1532: 1527: 1523: 1519: 1515: 1509: 1507: 1503: 1499: 1492: 1490: 1488: 1483: 1479: 1475: 1474:colour vision 1470: 1468: 1464: 1460: 1456: 1452: 1447: 1445: 1441: 1437: 1429: 1427: 1423: 1420: 1416: 1412: 1407: 1403: 1402:Colour vision 1395: 1393: 1389: 1388:) in humans. 1387: 1386:normal vision 1383: 1377: 1375: 1370: 1365: 1363: 1359: 1358: 1352: 1348: 1347:Visual acuity 1342: 1339:The eye of a 1337: 1331:Visual acuity 1330: 1325: 1323: 1320: 1316: 1312: 1309: 1304: 1301: 1295: 1291: 1288: 1279: 1277: 1275: 1271: 1267: 1261: 1259: 1255: 1249: 1247: 1243: 1237: 1234: 1229: 1227: 1223: 1219: 1215: 1211: 1206: 1203: 1199: 1195: 1188: 1183: 1178: 1170: 1168: 1166: 1162: 1157: 1152: 1149: 1141: 1139: 1137: 1133: 1129: 1128: 1122: 1120: 1116: 1115: 1109: 1107: 1103: 1097: 1095: 1092:) absorb the 1091: 1087: 1083: 1078: 1074: 1070: 1066: 1061: 1059: 1055: 1051: 1043: 1041: 1035: 1033: 1031: 1027: 1023: 1019: 1015: 1011: 1007: 998: 995: 992: 991: 990: 984: 982: 980: 974: 972: 971: 966: 962: 958: 950: 948: 944: 942: 941:phased arrays 938: 934: 930: 922: 917: 909: 903: 898: 893: 892:Arthropod eye 887: 880:Compound eyes 879: 877: 875: 872: 868: 863: 861: 860: 855: 851: 847: 842: 835: 833: 831: 827: 823: 815: 811: 807: 803: 799: 797: 796:Luneburg lens 793: 789: 781: 776: 769: 767: 763: 761: 760: 755: 754: 750: 742: 740: 737: 732: 728: 726: 723:, which have 722: 718: 714: 710: 706: 701: 699: 698:box jellyfish 694: 692: 687: 679: 677: 675: 671: 667: 663: 659: 655: 651: 649: 641: 639: 637: 633: 629: 625: 621: 613: 611: 609: 605: 600: 598: 594: 590: 586: 581: 579: 575: 571: 567: 563: 559: 555: 554:compound eyes 546: 544: 542: 538: 534: 530: 526: 522: 518: 514: 510: 506: 502: 498: 497:field of view 494: 489: 486: 482: 478: 476: 475:hyperspectral 472: 471:mantis shrimp 464: 463: 457: 453: 451: 446: 441: 439: 435: 431: 427: 423: 419: 415: 411: 407: 403: 399: 395: 391: 387: 383: 380:known as the 379: 375: 371: 367: 363: 357: 351: 349: 346:, which have 345: 341: 337: 333: 329: 325: 317: 313: 309: 303: 298: 291: 289: 287: 283: 279: 275: 271: 266: 262: 260: 256: 252: 251:compound eyes 247: 245: 244:visual cortex 241: 237: 233: 229: 225: 221: 217: 212: 210: 209:visual system 206: 202: 198: 194: 190: 189:sensory organ 186: 174: 168: 164: 161: 157: 154: 152: 148: 145: 141: 138: 136: 132: 129: 126: 124: 120: 115: 112: 109: 107: 103: 98: 94: 91:Diagram of a 88: 83: 79: 75: 70: 65: 60: 57: 53: 49: 45: 38: 34: 33:Eye (cyclone) 30: 26: 22: 4628: 4522:Ciliary body 4362:Diencephalon 4361: 4346:Parasol cell 4330:Bipolar cell 4181:Pars plicata 4163:Ciliary body 4031: 3896:the original 3891: 3859:Plenum Press 3857:. New York: 3854: 3847:Bibliography 3801: 3797: 3743: 3739: 3685: 3679: 3653:. Retrieved 3628: 3624: 3611: 3586: 3578: 3566: 3555:. Retrieved 3538: 3534: 3524: 3513:. Retrieved 3485: 3478: 3467:. Retrieved 3458: 3449: 3438:. Retrieved 3410: 3403: 3395: 3389:. Retrieved 3361: 3354: 3349:, p. 28 3342: 3309: 3303: 3278: 3274: 3249: 3220:. Retrieved 3216:the original 3206: 3165: 3159: 3153: 3118: 3112: 3091: 3086: 3041: 3037: 3027: 2994: 2990: 2986: 2982: 2976: 2951: 2947: 2941: 2922: 2916: 2904: 2859: 2855: 2845: 2820: 2816: 2809: 2798:. Retrieved 2770: 2764: 2755: 2719: 2715: 2709: 2697:. Retrieved 2690:the original 2681: 2674: 2647: 2643: 2633: 2600: 2596: 2590: 2565: 2561: 2557: 2551: 2532: 2528: 2518: 2506:. Retrieved 2499:the original 2478: 2474: 2461: 2450:the original 2437: 2424: 2381: 2377: 2367: 2342: 2338: 2284: 2280: 2270: 2259: 2248: 2223: 2217: 2211: 2199:. Retrieved 2185: 2144: 2140: 2134: 2117: 2113: 2107: 2062: 2058: 2045: 2026: 2022: 1980: 1976: 1972: 1966: 1954:. Retrieved 1941: 1932: 1924:the original 1913: 1902:. Retrieved 1882: 1875: 1864:. Retrieved 1836: 1829: 1804: 1796: 1763: 1757: 1667: 1645: 1616:Eye movement 1521: 1513: 1510: 1500: 1496: 1493:Pigmentation 1471: 1448: 1433: 1424: 1408: 1405: 1390: 1378: 1366: 1354: 1345: 1321: 1317: 1313: 1296: 1292: 1283: 1262: 1250: 1238: 1230: 1210:bird of prey 1207: 1191: 1155: 1153: 1148:ciliary body 1147: 1145: 1132:brittle star 1130:, a type of 1125: 1124:The body of 1123: 1112: 1110: 1105: 1098: 1090:optical axes 1062: 1053: 1050:Strepsiptera 1047: 1039: 1002: 988: 975: 968: 954: 945: 933:polarisation 926: 886:Compound eye 864: 857: 843: 839: 819: 800: 785: 764: 757: 751: 746: 733: 729: 702: 695: 683: 674:ion channels 672:but TRP are 652: 645: 617: 601: 582: 550: 527:. They have 509:caterpillars 490: 479: 468: 460: 450:Strepsiptera 442: 352: 321: 267: 263: 248: 213: 195:to perceive 184: 182: 144:A01.1.00.007 140:A15.2.00.001 127: 74:Compound eye 56: 4614:Eye disease 4594:Keratocytes 4483:Conjunctiva 4440:Ora serrata 4378:Muller glia 4342:Midget cell 4062:Endothelium 4052:Dua's layer 3919:Adobe Flash 2911:, p. 8 2699:13 November 2481:: 147–177. 2253:Animal Eyes 1606:Eye disease 1487:optic nerve 1355:cycles per 1308:Glyptonotus 1218:ultraviolet 1187:mollusc eye 1082:pseudopupil 1069:dragonflies 993:refracting, 947:ommatidia. 937:diffraction 717:cephalopods 632:crustaceans 624:cephalopods 620:vertebrates 593:entrainment 566:rhabdomeric 422:cephalopods 402:transparent 398:optic nerve 370:vertebrates 259:crustaceans 240:optic nerve 230:to form an 117:Identifiers 4649:Categories 4455:of the eye 4428:Optic disc 4186:Pars plana 4057:Descemet's 4037:Epithelium 3655:2008-09-16 3587:The Senses 3557:2012-01-01 3535:J Exp Biol 3515:2020-10-19 3469:2015-06-03 3440:2020-10-19 3391:2020-10-19 3222:2012-09-01 2987:Drosophila 2800:2008-09-15 2568:(2): 137. 2562:Alcheringa 2219:BioScience 1904:2020-10-19 1866:2020-10-19 1837:Psychology 1679:References 1611:Eye injury 1576:(focusing) 1529:r-opsins. 1467:blind spot 1415:cone cells 1351:cone cells 1326:Physiology 1258:crystallin 1058:trilobites 979:ommatidium 929:ommatidium 705:gastropods 654:Pit vipers 574:gastropods 481:Trilobites 445:arthropods 430:amphibians 386:cone cells 368:. 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