Eye - Knowledge

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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).

1375:, 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. 1568: 661:, 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. 1417:"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. 4088: 98: 1556: 817: 1424:, 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 308: 927: 919: 467: 1193: 319: 786: 1305:

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.

1259:, 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. 80: 1119:. 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. 1487:. 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 1015:

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).

908: 805:, 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 942:, 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 1267:, 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 1495:

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

880:, 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 699:

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.

1544:, 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. 1071:. 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. 1211:) 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 809:), 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. 776:

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

984:, 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.) 1308:

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.

1500:. 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. 1300:

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.

575:, 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 1274:

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.

1149:. 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. 4125:

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

954:). 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. 741:

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.

1519:, 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. 2123:

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".

463:, 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. 1235:, 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 1333:

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.

1174:), 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 738:

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".

1960: 375:. 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 1287:. This increased refractive power and again eased circulatory problems. Formation of a nontransparent ring allowed more blood vessels, more circulation, and larger eye sizes. 530:

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

843:, 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. 1930: 2441: 2725:

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

1468:) 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 ( 773:, has two lenses in each eye, arranged like those in a telescope. Such arrangements are rare and poorly understood, but represent an alternative construction. 3319:

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,

4106: 2274: 272:. 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. 2205: 4111: 2061:

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

1239:, but this is not the case, as the reversed roles of their respective ciliary and rhabdomeric opsin classes and different lens crystallins show. 4107: 4112: 4092: 3898: 3968: 3877: 3611: 3505: 3430: 3381: 2941: 1902: 1856: 1829: 154: 150: 3473: 1948: 672: 4098: 1555: 3519: 3444: 3395: 1870: 2776: 1908: 873:

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.

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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)

610:. These are not considered eyes because they lack enough structure to be considered an organ, and do not produce an image. 431:, regulating the amount of light that enters the eye and reducing aberrations when there is enough light. The eyes of most 4094: 2460: 1769: 704:. Such a non-homogeneous lens is necessary for the focal length to drop from about 4 times the lens radius, to 2.5 radii. 613:

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?".

1492: 4512: 4481: 4473: 4153: 3124: 1651: 790: 603: 4113: 4104: 4101: 4308: 4298: 680: 668: 4110: 4109: 403:(for low-light contrasts) in the retina detect and convert light into neural signals which are transmitted to the 4093: 3961: 1631: 1019:, the parabolic surfaces of the inside of each facet focus light from a reflector to a sensor array. Long-bodied 2509: 4116: 3990: 1601: 827:

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

4670: 4303: 4259: 4117: 2536:"Optics and phylogeny: is there an insight? The evolution of superposition eyes in the Decapoda (Crustacea)" 1318: 349:. In other organisms, particularly prey animals, eyes are located to maximise the field of view, such as in 288: 62: 2201: 1067:

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

296: 177: 161: 145: 671:). However, pit organs are fitted with receptors rather different from photoreceptors, namely a specific 4619: 4424: 4348: 4286: 4274: 4229: 1488: 1243: 412: 58: 4119: 3043:

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

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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

1360:, or resolving power, is "the ability to distinguish fine detail" and is the property of 1219:

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".

2456: 4659: 4572: 4507: 4344: 3749:

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

3603: 3079: 3044: 2970: 2136: 1999: 1934: 1641: 1412: 1396: 1357: 1204: 902: 806: 798: 708: 564: 507: 485: 481: 254: 219: 199: 43: 3846: 3778: 3728: 3344: 3305: 2426: 2329: 2187: 2030:"Exceptional Variation on a Common Theme: the Evolution of Crustacean Compound Eyes" 1472:) intensities. Rods are distributed throughout the retina but there are none at the 459:

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

27:

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".

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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.

664: 642: 634: 630: 432: 380: 269: 3925: 3596: 3515: 3391: 3120:"Evolution of the vertebrate eye: opsins, photoreceptors, retina and eye cup" 2700: 1790: 170: 4414: 4409: 4328: 3982: 3440: 3021: 2887: 2171: 2090: 1866: 1606: 1454: 1425: 1421: 1379: 1224: 1208: 1129: 1124: 1096: 1079: 967: 735: 614: 584: 491: 455: 440: 396: 387:, the eye allows light to enter and project onto a light-sensitive layer of 322: 166: 103: 31: 17: 3838: 3770: 3336: 3155: 3069: 2906: 2789: 2676: 2635: 2505: 2418: 2321: 2179: 2109: 2063:"Assembly of the cnidarian camera-type eye from vertebrate-like components" 714:

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

215: 3647: 534:, some of the simplest eyes, are found in animals such as some of the 4579: 4489: 4243: 4029: 4001: 3192: 1515:

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

392: 376: 366: 350: 334: 326: 265: 226: 3137: 2385:"A Novel Vertebrate Eye Using Both Refractive and Reflective Optics" 2242: 1460:

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

907: 484:

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

53:"Eyeball", "Eyes", and "Ocular" redirect here. For other uses, see 4219: 1663: 1512: 1345: 1191: 1112: 1028: 925: 917: 906: 836: 815: 784: 676: 535: 515: 465: 444: 428: 404: 354: 317: 306: 246: 242: 211: 183: 133: 39: 3935: 2651:"Dimensional limits for arthropod eyes with superposition optics" 4133: 1212: 758:

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

567:

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

436: 3950: 2765:"Variations in the optical properties of the compound eyes of 1931:"What animal has a more sophisticated eye, Octopus or Insect?" 1573:

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.

476:) are considered the most complex in the whole animal kingdom. 3422:

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

1527:

cells of vertebrates express r-opsins, suggesting that their

1283:. Separation of the forward layer again formed a humour, the 4087: 3936:

Webvision. The organisation of the retina and visual system.

3118:

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

2863:"Unique system of photoreceptors in sea urchin tube feet" 35: 1496:

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

526:) which usually provides only a rough image, but (as in 2227:

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

1128:. This type of eye consists of a cluster of numerous 249:

through neural pathways that connect the eye via the

4597: 4558: 4472: 4463: 4397: 4317: 4252: 4242: 4205: 4172: 4144: 4132: 4028: 4000: 3989: 3245: 3243: 1326:increase the amount of light that can be captured. 160: 144: 132: 127: 115: 110: 72: 3899:"Inside the Eye: Nature's Most Exquisite Creation" 3595: 2204:. National Institute of General Medical Sciences. 1813: 214:and converts it into electro-chemical impulses in 3602:. Cambridge: Cambridge University Press. p. 3251:The Evolution of Eyes: Where Do Lenses Come From? 679:. The main difference is that photoreceptors are 506:have one pair of large simple eyes with a narrow 427:. Muscles around the iris change the size of the 3926:Anatomy of the eye – flash animated interactive. 3684: 3682: 3680: 3678: 3676: 4091: 2867:Proceedings of the National Academy of Sciences 2288:"A Chiton Uses Aragonite Lenses to Form Images" 2275:Perceiving in Depth, Volume 1: Basic Mechanisms 494:, now extinct, had unique compound eyes. Clear 3798: 3796: 3794: 3792: 3790: 3788: 3744: 3742: 3740: 3738: 2440:Völkel, R; Eisner, M; Weible, KJ (June 2003). 1523:cells in the eye with r-opsins. However, some 3962: 2343: 2341: 2339: 1820:. New York: Oxford University Press. p. 1162:constitutes the cells of the dilator muscle. 579:. These two groups are not monophyletic; the 8: 2023: 2021: 2019: 2017: 1391:pair, equivalent to a 1 arcminute gap in an 510:, augmented by an array of smaller eyes for 3594:Barlow, Horace Basil; Mollon, J.D. (1982). 3279: 3277: 3275: 3273: 3271: 3269: 1762: 1760: 1758: 1756: 1754: 1752: 1750: 1748: 1746: 1744: 1742: 1740: 1738: 1736: 1734: 1732: 1730: 1728: 1726: 1724: 364:The first proto-eyes evolved among animals 4469: 4249: 4141: 3997: 3969: 3955: 3947: 3542:"The size of ommatidia in apposition eyes" 3321:Proceedings of the Royal Society of London 1722: 1720: 1718: 1716: 1714: 1712: 1710: 1708: 1706: 1704: 1099:which one observes "head-on" (along their 291:to effect circadian adjustment and to the 225:In higher organisms, the eye is a complex 96: 78: 3828: 3145: 3078: 3068: 2896: 2886: 2666: 2551: 2408: 2311: 2099: 2089: 2045: 3864:Ali, Mohamed Ather; Klyne, M.A. (1985). 3581: 3357: 3225:. Library.thinkquest.org. Archived from 2919: 2763:Jochen Zeil; Maha M. Al-Mutairi (1996). 922:Anatomy of the compound eye of an insect 218:(neurones). It is part of an organism's 3113: 3111: 2758: 2756: 1816:Blindspots: The Many Ways We Cannot See 1700: 1675: 1551: 867:, also use reflector eyes. The scallop 583:also possess ciliated cells, and some 568: 550:, compound eyes are adapted to see the 2699:(PhD). Lund University. Archived from 1223:has much greater visual acuity than a 1107:, while those to one side reflect it. 876:There is at least one vertebrate, the 181: 69: 3805:"Eye Evolution: the Blurry Beginning" 1933:. BioMedia Associates. Archived from 571:can achieve greater sensitivity than 237:it through an adjustable assembly of 7: 3803:Nilsson, E.; Arendt, D. (Dec 2008). 2690:Greiner, Birgit (16 December 2005). 1929:Kirk, Molly; Denning, David (2001). 673:transient receptor potential channel 333:Complex eyes distinguish shapes and 3286:Journal of Comparative Physiology A 2777:The Journal of Experimental Biology 2608:Arthropod Structure and Development 2028:Cronin, T.W.; Porter, M.L. (2008). 1783:10.1146/annurev.ne.15.030192.000245 2208:from the original on 13 March 2020 1963:from the original on 9 August 2010 1561:The structures of the eye labelled 470:The eyes of a mantis shrimp (here 25: 3105:. Oxford: Oxford University Press 3045:"Squid Pax-6 and eye development" 2649:Meyer-Rochow, VB; Gal, J (2004). 2350:Journal of Comparative Physiology 2034:Evolution: Education and Outreach 1851:. Belmont: Wadsworth Publishing. 1291:Relationship to life requirements 988:absorbed by the dark wall of the 2137:10.1111/j.1463-6395.2006.00216.x 1897:. Psychology Press. p. 20. 1566: 1554: 1428:, from which the more sensitive 3692:The Quarterly Review of Biology 3662:from the original on 2012-11-20 3564:from the original on 2016-08-31 3522:from the original on 2023-01-17 3476:from the original on 2015-05-04 3447:from the original on 2023-01-17 3398:from the original on 2023-01-17 2807:from the original on 2009-02-25 1911:from the original on 2023-01-17 1873:from the original on 2023-01-17 1227:, and in some cases can detect 42:. For region of a cyclone, see 2442:"Miniaturized imaging systems" 2202:"Circadian Rhythms Fact Sheet" 1585:Accommodation (vertebrate eye) 835:which wipe the eye and spread 602:, which is sufficient for the 257:and other areas of the brain. 1: 3636:Brain, Behavior and Evolution 3470:hyperphysics.phy-astr.gsu.edu 3466:"The Retina of the Human Eye" 3373:The Image Processing Handbook 2498:10.1146/annurev.ento.42.1.147 2457:10.1016/S0167-9317(03)00102-3 1770:Annual Review of Neuroscience 1091:at the observer, is called a 855:Many small organisms such as 546:. In organisms dwelling near 522:, have a type of simple eye ( 281:photosensitive ganglion cells 4615:Optical coherence tomography 4369:Photosensitive ganglion cell 3627:Fernald, Russell D. (1997). 3500:. McGraw-Hill Professional. 3425:. McGraw-Hill Professional. 3419:Klaassen, Curtis D. (2001). 3249:Fernald, Russell D. (2001). 2971:10.1016/0959-437X(95)80029-8 2668:10.1016/j.visres.2004.04.009 2567:Ritchie, Alexander (1985). " 2000:10.1016/0022-1910(74)90087-0 1988:Journal of Insect Physiology 1957:National Wildlife Federation 1420:The most sensitive pigment, 1082:, have specialised zones of 913:scanning electron microscope 4365:Giant retina ganglion cells 4154:Capillary lamina of choroid 3125:Nature Reviews Neuroscience 2486:Annual Review of Entomology 2449:Microelectronic Engineering 1949:"Who You Callin' "Shrimp"?" 1652:Simple eye in invertebrates 791:anterior chamber of the eye 681:G-protein coupled receptors 4692: 4309:Retinal pigment epithelium 4299:External limiting membrane 3101:Conway-Morris, S. (1998). 3049:Proc. Natl. Acad. Sci. USA 2479:"Visual Acuity in Insects" 1953:National Wildlife Magazine 1812:Breitmeyer, Bruno (2010). 1483:Cones are responsible for 1410: 1364:. It is often measured in 1185: 1095:. This occurs because the 900: 894: 669:infrared sensing in snakes 52: 29: 4085: 3830:10.1016/j.cub.2008.10.025 3323:. B 215 (1201): 433–450. 2840:10.1007/s00435-003-0085-0 2747:10.1080/08912969709386565 2628:10.1016/j.asd.2006.06.003 2593:10.1080/03115518508618961 2553:10.1163/18759866-06704001 2410:10.1016/j.cub.2008.11.061 2313:10.1016/j.cub.2011.03.033 2047:10.1007/s12052-008-0085-0 1632:Lens (vertebrate anatomy) 1371:(CPD), which measures an 1117:Dioptromysis paucispinosa 454:The compound eyes of the 176: 95: 77: 3103:The Crucible of Creation 2540:Contributions to Zoology 1602:Emission theory (vision) 1395:, corresponds to 20/20 ( 930:Arthropods such as this 657:Pit eyes, also known as 473:Odontodactylus scyllarus 285:retinohypothalamic tract 210:information. It detects 55:Eyeball (disambiguation) 4304:Layer of rods and cones 4260:Inner limiting membrane 3629:"The Evolution of Eyes" 3263:64: "The Eye in Focus". 3022:10.1126/science.7892602 2888:10.1073/pnas.1018495108 2172:10.1126/science.1127889 2091:10.1073/pnas.0800388105 1047:Parabolic superposition 1010:parabolic superposition 369: million years ago 283:send signals along the 63:Ocular (disambiguation) 4126: 3370:Russ, John C. (2006). 3337:10.1098/rspb.1982.0052 3070:10.1073/pnas.94.6.2421 2959:Curr. Opin. Genet. Dev 2790:10.1242/jeb.199.7.1569 2534:Gaten, Edward (1998). 2477:Land, Michael (1997). 2451:. 67–68 (1): 461–472. 1845:Nairne, James (2005). 1354: 1200: 935: 923: 915: 828: 794: 675:(TRP channels) called 477: 371:about the time of the 330: 315: 297:pupillary light reflex 289:suprachiasmatic nuclei 178:Anatomical terminology 46:. For other uses, see 34:. For the letter, see 4620:Eye care professional 4425:Foveal avascular zone 4287:Outer plexiform layer 4275:Inner plexiform layer 4230:Iris sphincter muscle 4123: 3866:Vision in Vertebrates 3540:Barlow, H.B. (1952). 3497:Optical System Design 2569:Ainiktozoon loganense 1449:used for vision: the 1349: 1244:photoreceptor protein 1195: 929: 921: 910: 901:Further information: 819: 788: 469: 419:, possess a focusing 399:(for colour) and the 321: 310: 59:Eyes (disambiguation) 38:. For the pupil, see 4640:Physiological Optics 4610:Ocular immune system 4349:Retina ganglion cell 3921:Evolution of the eye 3909:on January 14, 2016. 3757:(11–12): 1151–1162. 3582:Ali & Klyne 1985 3558:10.1242/jeb.29.4.667 3358:Ali & Klyne 1985 2920:Ali & Klyne 1985 2767:Uca lactea annulipes 1937:on 26 February 2017. 1637:Nictitating membrane 1188:Evolution of the eye 799:eyes of most mammals 702:spherical aberration 594:Some organisms have 417:vitreous humour 48:Eye (disambiguation) 4464:Anatomical regions 4325:Photoreceptor cells 4292:Outer nuclear layer 4280:Inner nuclear layer 4270:Ganglion cell layer 4225:Iris dilator muscle 4020:Trabecular meshwork 3903:National Geographic 3897:(14 January 2016). 3821:2008CBio...18R1096N 3815:(23): R1096–R1098. 3329:1982RSPSB.215..433M 3185:1988Natur.336...86T 3061:1997PNAS...94.2421T 3014:1995Sci...267.1788H 3008:(5205): 1788–1792. 2879:2011PNAS..108.8367U 2739:1997HBio...12..229H 2620:2006ArtSD..35..231M 2585:1985Alch....9..117R 2515:on 23 November 2004 2401:2009CBio...19..108W 2304:2011CBio...21..665S 2164:2006Sci...313.1914F 2158:(5795): 1914–1918. 2082:2008PNAS..105.8989K 1447:photoreceptor cells 1043:instead of lenses. 1021:decapod crustaceans 934:have compound eyes. 4127: 3763:10.1002/bies.20828 3298:10.1007/BF00616747 3256:2006-03-19 at the 2727:Historical Biology 2706:on 9 February 2013 2362:10.1007/BF00661380 1959:. 1 October 2010. 1373:angular resolution 1355: 1233:parallel evolution 1203:Photoreception is 1201: 1176:mucopolysaccharide 996:Superposition eyes 936: 924: 916: 829: 795: 730:, and once in the 691:Spherical lens eye 569:superposition eyes 478: 373:Cambrian explosion 331: 316: 4653: 4652: 4645:Visual perception 4593: 4592: 4560:Posterior segment 4528:Posterior chamber 4459: 4458: 4361:Bistratified cell 4265:Nerve fiber layer 4238: 4237: 4182:Ciliary processes 4083: 4082: 3879:978-0-306-42065-8 3648:10.1159/000113339 3613:978-0-521-24474-9 3507:978-0-07-134916-1 3432:978-0-07-134721-1 3383:978-0-8493-7254-4 2943:978-3-540-08837-0 2873:(20): 8367–8372. 2661:(19): 2213–2223. 2076:(26): 8989–8993. 1904:978-0-86377-450-8 1858:978-0-495-03150-5 1831:978-0-19-539426-9 1548:Additional images 1407:Colour perception 1403:vertebrate eyes. 1196:Evolution of the 1138:Ophiocoma wendtii 1086:organised into a 781:Refractive cornea 625:Non-compound eyes 608:circadian rhythms 512:peripheral vision 192: 191: 187: 16:(Redirected from 4683: 4630:Refractive error 4568:Vitreous chamber 4513:Anterior chamber 4474:Anterior segment 4470: 4250: 4159:Bruch's membrane 4142: 4134:Uvea / vascular 4090: 4010:Episcleral layer 3998: 3971: 3964: 3957: 3948: 3910: 3905:. Archived from 3883: 3851: 3850: 3832: 3800: 3783: 3782: 3746: 3733: 3732: 3686: 3671: 3670: 3668: 3667: 3633: 3624: 3618: 3617: 3601: 3591: 3585: 3579: 3573: 3572: 3570: 3569: 3537: 3531: 3530: 3528: 3527: 3491: 3485: 3484: 3482: 3481: 3462: 3456: 3455: 3453: 3452: 3416: 3410: 3409: 3404: 3403: 3367: 3361: 3355: 3349: 3348: 3316: 3310: 3309: 3281: 3264: 3247: 3238: 3237: 3235: 3234: 3223:"Eye-Evolution?" 3219: 3213: 3212: 3193:10.1038/336086a0 3166: 3160: 3159: 3149: 3115: 3106: 3099: 3093: 3092: 3082: 3072: 3055:(6): 2421–2426. 3040: 3034: 3033: 2989: 2983: 2982: 2954: 2948: 2947: 2929: 2923: 2917: 2911: 2910: 2900: 2890: 2858: 2852: 2851: 2822: 2816: 2815: 2813: 2812: 2806: 2784:(7): 1569–1577. 2773: 2760: 2751: 2750: 2733:(3–4): 229–263. 2722: 2716: 2715: 2713: 2711: 2705: 2698: 2687: 2681: 2680: 2670: 2646: 2640: 2639: 2603: 2597: 2596: 2564: 2558: 2557: 2555: 2531: 2525: 2524: 2522: 2520: 2514: 2508:. Archived from 2483: 2474: 2468: 2467: 2465: 2459:. Archived from 2446: 2437: 2431: 2430: 2412: 2380: 2374: 2373: 2345: 2334: 2333: 2315: 2283: 2277: 2272: 2266: 2261: 2255: 2254: 2224: 2218: 2217: 2215: 2213: 2198: 2192: 2191: 2147: 2141: 2140: 2120: 2114: 2113: 2103: 2093: 2067: 2066:(Free full text) 2058: 2052: 2051: 2049: 2025: 2012: 2011: 1994:(8): 1565–1591. 1979: 1973: 1972: 1970: 1968: 1945: 1939: 1938: 1926: 1920: 1919: 1917: 1916: 1888: 1882: 1881: 1879: 1878: 1842: 1836: 1835: 1819: 1809: 1803: 1802: 1764: 1683: 1680: 1591:Adaptation (eye) 1570: 1558: 1476:and none at the 1265:refractive index 1205:phylogenetically 697:refractive index 370: 359:monocular vision 347:depth perception 343:binocular vision 184:edit on Wikidata 100: 82: 70: 21: 4691: 4690: 4686: 4685: 4684: 4682: 4681: 4680: 4656: 4655: 4654: 4649: 4589: 4554: 4543:Capsule of lens 4498:Lacrimal system 4465: 4455: 4415:Parafoveal area 4410:Perifoveal area 4393: 4337:Horizontal cell 4313: 4234: 4201: 4168: 4164:Sattler's layer 4135: 4128: 4122: 4079: 4024: 4015:Schlemm's canal 3993: 3985: 3977:Anatomy of the 3975: 3917: 3893: 3890: 3888:Further reading 3880: 3863: 3860: 3855: 3854: 3809:Current Biology 3802: 3801: 3786: 3748: 3747: 3736: 3688: 3687: 3674: 3665: 3663: 3631: 3626: 3625: 3621: 3614: 3593: 3592: 3588: 3580: 3576: 3567: 3565: 3539: 3538: 3534: 3525: 3523: 3508: 3493: 3492: 3488: 3479: 3477: 3464: 3463: 3459: 3450: 3448: 3433: 3418: 3417: 3413: 3401: 3399: 3384: 3369: 3368: 3364: 3356: 3352: 3318: 3317: 3313: 3283: 3282: 3267: 3258:Wayback Machine 3248: 3241: 3232: 3230: 3221: 3220: 3216: 3179:(6194): 86–88. 3168: 3167: 3163: 3138:10.1038/nrn2283 3132:(12): 960–976. 3117: 3116: 3109: 3100: 3096: 3042: 3041: 3037: 2991: 2990: 2986: 2956: 2955: 2951: 2944: 2931: 2930: 2926: 2918: 2914: 2860: 2859: 2855: 2824: 2823: 2819: 2810: 2808: 2804: 2771: 2762: 2761: 2754: 2724: 2723: 2719: 2709: 2707: 2703: 2696: 2689: 2688: 2684: 2655:Vision Research 2648: 2647: 2643: 2605: 2604: 2600: 2566: 2565: 2561: 2533: 2532: 2528: 2518: 2516: 2512: 2481: 2476: 2475: 2471: 2463: 2444: 2439: 2438: 2434: 2389:Current Biology 2382: 2381: 2377: 2347: 2346: 2337: 2292:Current Biology 2285: 2284: 2280: 2273: 2269: 2262: 2258: 2243:10.2307/1311112 2226: 2225: 2221: 2211: 2209: 2200: 2199: 2195: 2149: 2148: 2144: 2122: 2121: 2117: 2065: 2060: 2059: 2055: 2027: 2026: 2015: 1981: 1980: 1976: 1966: 1964: 1947: 1946: 1942: 1928: 1927: 1923: 1914: 1912: 1905: 1890: 1889: 1885: 1876: 1874: 1859: 1844: 1843: 1839: 1832: 1811: 1810: 1806: 1766: 1765: 1702: 1697: 1692: 1687: 1686: 1681: 1677: 1672: 1658:Tapetum lucidum 1647:Orbit (anatomy) 1612:Eye development 1597:Capsule of lens 1581: 1574: 1571: 1562: 1559: 1550: 1506: 1466:black-and-white 1443: 1415: 1409: 1352:red-tailed hawk 1344: 1339: 1293: 1190: 1184: 1172:hyaluronic acid 1155: 1057: 1049: 998: 964: 962:Apposition eyes 932:blue bottle fly 905: 899: 893: 859:, copepods and 849: 783: 756: 754:Multiple lenses 693: 655: 627: 573:apposition eyes 560: 552:infra-red light 504:Jumping spiders 488:colour vision. 423:, and often an 365: 325:, a refractive 305: 295:to control the 202:that allows an 188: 153: 106: 91: 89:Antarctic krill 66: 51: 28: 23: 22: 15: 12: 11: 5: 4689: 4687: 4679: 4678: 4673: 4671:Sensory organs 4668: 4658: 4657: 4651: 4650: 4648: 4647: 4642: 4637: 4632: 4627: 4622: 4617: 4612: 4607: 4601: 4599: 4595: 4594: 4591: 4590: 4588: 4587: 4582: 4577: 4576: 4575: 4564: 4562: 4556: 4555: 4553: 4552: 4551: 4550: 4548:Zonule of Zinn 4545: 4535: 4530: 4525: 4520: 4518:Aqueous humour 4515: 4510: 4505: 4478: 4476: 4467: 4461: 4460: 4457: 4456: 4454: 4453: 4448: 4447: 4446: 4436: 4435: 4434: 4433: 4432: 4427: 4417: 4412: 4401: 4399: 4395: 4394: 4392: 4391: 4321: 4319: 4315: 4314: 4312: 4311: 4306: 4301: 4295: 4294: 4289: 4283: 4282: 4277: 4272: 4267: 4262: 4256: 4254: 4247: 4240: 4239: 4236: 4235: 4233: 4232: 4227: 4222: 4217: 4211: 4209: 4203: 4202: 4200: 4199: 4194: 4189: 4187:Ciliary muscle 4184: 4178: 4176: 4170: 4169: 4167: 4166: 4161: 4156: 4150: 4148: 4139: 4130: 4129: 4086: 4084: 4081: 4080: 4078: 4077: 4076: 4075: 4070: 4065: 4060: 4055: 4050: 4040: 4034: 4032: 4026: 4025: 4023: 4022: 4017: 4012: 4006: 4004: 3995: 3987: 3986: 3976: 3974: 3973: 3966: 3959: 3951: 3945: 3944: 3939: 3933: 3923: 3916: 3915:External links 3913: 3912: 3911: 3889: 3886: 3885: 3884: 3878: 3859: 3856: 3853: 3852: 3784: 3734: 3705:10.1086/416840 3699:(3): 281–322. 3672: 3642:(4): 253–259. 3619: 3612: 3586: 3574: 3552:(4): 667–674. 3532: 3506: 3486: 3457: 3431: 3411: 3382: 3362: 3350: 3311: 3292:(6): 751–762. 3265: 3261:Karger Gazette 3239: 3214: 3161: 3107: 3094: 3035: 2984: 2965:(5): 602–609. 2949: 2942: 2924: 2912: 2853: 2834:(4): 191–209. 2817: 2752: 2717: 2682: 2641: 2614:(4): 231–245. 2598: 2559: 2546:(4): 223–236. 2526: 2469: 2466:on 2008-10-01. 2432: 2395:(2): 108–114. 2375: 2356:(4): 297–316. 2335: 2298:(8): 665–670. 2278: 2267: 2256: 2237:(5): 298–307. 2219: 2193: 2142: 2125:Acta Zoologica 2115: 2053: 2040:(4): 463–475. 2013: 1974: 1940: 1921: 1903: 1883: 1857: 1837: 1830: 1804: 1699: 1698: 1696: 1693: 1691: 1688: 1685: 1684: 1674: 1673: 1671: 1668: 1667: 1666: 1661: 1654: 1649: 1644: 1639: 1634: 1629: 1624: 1619: 1614: 1609: 1604: 1599: 1594: 1593:(night vision) 1588: 1580: 1577: 1576: 1575: 1572: 1565: 1563: 1560: 1553: 1549: 1546: 1505: 1502: 1464:) monochrome ( 1442: 1441:Rods and cones 1439: 1411:Main article: 1408: 1405: 1343: 1340: 1338: 1335: 1292: 1289: 1285:aqueous humour 1253:pinhole camera 1237:cephalopod eye 1186:Main article: 1183: 1180: 1154: 1151: 1105:incident light 1076:praying mantis 1056: 1053: 1048: 1045: 1012: 1011: 1008: 1007:reflecting and 1005: 997: 994: 963: 960: 895:Main article: 892: 889: 848: 847:Reflector eyes 845: 782: 779: 755: 752: 726:, once in the 722:, once in the 718:, once in the 692: 689: 654: 651: 626: 623: 619:Fresnel lenses 596:photosensitive 591:possess both. 559: 556: 548:deep-sea vents 540:photosensitive 514:. Some insect 313:European bison 304: 301: 293:pretectal area 190: 189: 180: 174: 173: 164: 158: 157: 148: 142: 141: 136: 130: 129: 125: 124: 119: 113: 112: 108: 107: 101: 93: 92: 83: 75: 74: 26: 24: 14: 13: 10: 9: 6: 4: 3: 2: 4688: 4677: 4676:Visual system 4674: 4672: 4669: 4667: 4664: 4663: 4661: 4646: 4643: 4641: 4638: 4636: 4635:Accommodation 4633: 4631: 4628: 4626: 4623: 4621: 4618: 4616: 4613: 4611: 4608: 4606: 4603: 4602: 4600: 4596: 4586: 4583: 4581: 4578: 4574: 4573:Vitreous body 4571: 4570: 4569: 4566: 4565: 4563: 4561: 4557: 4549: 4546: 4544: 4541: 4540: 4539: 4536: 4534: 4531: 4529: 4526: 4524: 4521: 4519: 4516: 4514: 4511: 4509: 4508:Fibrous tunic 4506: 4503: 4499: 4495: 4491: 4487: 4483: 4480: 4479: 4477: 4475: 4471: 4468: 4462: 4452: 4449: 4445: 4442: 4441: 4440: 4437: 4431: 4428: 4426: 4423: 4422: 4421: 4418: 4416: 4413: 4411: 4408: 4407: 4406: 4403: 4402: 4400: 4396: 4390: 4386: 4382: 4378: 4374: 4370: 4366: 4362: 4358: 4354: 4350: 4346: 4345:Amacrine cell 4342: 4338: 4334: 4330: 4326: 4323: 4322: 4320: 4316: 4310: 4307: 4305: 4302: 4300: 4297: 4296: 4293: 4290: 4288: 4285: 4284: 4281: 4278: 4276: 4273: 4271: 4268: 4266: 4263: 4261: 4258: 4257: 4255: 4251: 4248: 4245: 4241: 4231: 4228: 4226: 4223: 4221: 4218: 4216: 4213: 4212: 4210: 4208: 4204: 4198: 4195: 4193: 4190: 4188: 4185: 4183: 4180: 4179: 4177: 4175: 4171: 4165: 4162: 4160: 4157: 4155: 4152: 4151: 4149: 4147: 4143: 4140: 4137: 4131: 4074: 4071: 4069: 4066: 4064: 4061: 4059: 4056: 4054: 4051: 4049: 4046: 4045: 4044: 4041: 4039: 4036: 4035: 4033: 4031: 4027: 4021: 4018: 4016: 4013: 4011: 4008: 4007: 4005: 4003: 3999: 3996: 3992: 3991:Fibrous tunic 3988: 3984: 3980: 3972: 3967: 3965: 3960: 3958: 3953: 3952: 3949: 3943: 3940: 3937: 3934: 3931: 3927: 3924: 3922: 3919: 3918: 3914: 3908: 3904: 3900: 3896: 3892: 3891: 3887: 3881: 3875: 3871: 3867: 3862: 3861: 3857: 3848: 3844: 3840: 3836: 3831: 3826: 3822: 3818: 3814: 3810: 3806: 3799: 3797: 3795: 3793: 3791: 3789: 3785: 3780: 3776: 3772: 3768: 3764: 3760: 3756: 3752: 3745: 3743: 3741: 3739: 3735: 3730: 3726: 3722: 3718: 3714: 3710: 3706: 3702: 3698: 3694: 3693: 3685: 3683: 3681: 3679: 3677: 3673: 3661: 3657: 3653: 3649: 3645: 3641: 3637: 3630: 3623: 3620: 3615: 3609: 3605: 3600: 3599: 3590: 3587: 3584:, p. 161 3583: 3578: 3575: 3563: 3559: 3555: 3551: 3547: 3543: 3536: 3533: 3521: 3517: 3513: 3509: 3503: 3499: 3498: 3490: 3487: 3475: 3471: 3467: 3461: 3458: 3446: 3442: 3438: 3434: 3428: 3424: 3423: 3415: 3412: 3408: 3397: 3393: 3389: 3385: 3379: 3376:. CRC Press. 3375: 3374: 3366: 3363: 3359: 3354: 3351: 3346: 3342: 3338: 3334: 3330: 3326: 3322: 3315: 3312: 3307: 3303: 3299: 3295: 3291: 3287: 3280: 3278: 3276: 3274: 3272: 3270: 3266: 3262: 3259: 3255: 3252: 3246: 3244: 3240: 3229:on 2012-09-15 3228: 3224: 3218: 3215: 3210: 3206: 3202: 3198: 3194: 3190: 3186: 3182: 3178: 3174: 3173: 3165: 3162: 3157: 3153: 3148: 3143: 3139: 3135: 3131: 3127: 3126: 3121: 3114: 3112: 3108: 3104: 3098: 3095: 3090: 3086: 3081: 3076: 3071: 3066: 3062: 3058: 3054: 3050: 3046: 3039: 3036: 3031: 3027: 3023: 3019: 3015: 3011: 3007: 3003: 2999: 2995: 2988: 2985: 2980: 2976: 2972: 2968: 2964: 2960: 2953: 2950: 2945: 2939: 2935: 2928: 2925: 2921: 2916: 2913: 2908: 2904: 2899: 2894: 2889: 2884: 2880: 2876: 2872: 2868: 2864: 2857: 2854: 2849: 2845: 2841: 2837: 2833: 2829: 2828:Zoomorphology 2821: 2818: 2803: 2799: 2795: 2791: 2787: 2783: 2779: 2778: 2770: 2768: 2759: 2757: 2753: 2748: 2744: 2740: 2736: 2732: 2728: 2721: 2718: 2702: 2695: 2694: 2686: 2683: 2678: 2674: 2669: 2664: 2660: 2656: 2652: 2645: 2642: 2637: 2633: 2629: 2625: 2621: 2617: 2613: 2609: 2602: 2599: 2594: 2590: 2586: 2582: 2578: 2574: 2570: 2563: 2560: 2554: 2549: 2545: 2541: 2537: 2530: 2527: 2511: 2507: 2503: 2499: 2495: 2491: 2487: 2480: 2473: 2470: 2462: 2458: 2454: 2450: 2443: 2436: 2433: 2428: 2424: 2420: 2416: 2411: 2406: 2402: 2398: 2394: 2390: 2386: 2379: 2376: 2371: 2367: 2363: 2359: 2355: 2351: 2344: 2342: 2340: 2336: 2331: 2327: 2323: 2319: 2314: 2309: 2305: 2301: 2297: 2293: 2289: 2282: 2279: 2276: 2271: 2268: 2265: 2260: 2257: 2252: 2248: 2244: 2240: 2236: 2232: 2231: 2223: 2220: 2207: 2203: 2197: 2194: 2189: 2185: 2181: 2177: 2173: 2169: 2165: 2161: 2157: 2153: 2146: 2143: 2138: 2134: 2130: 2126: 2119: 2116: 2111: 2107: 2102: 2097: 2092: 2087: 2083: 2079: 2075: 2071: 2064: 2057: 2054: 2048: 2043: 2039: 2035: 2031: 2024: 2022: 2020: 2018: 2014: 2009: 2005: 2001: 1997: 1993: 1989: 1985: 1978: 1975: 1962: 1958: 1954: 1950: 1944: 1941: 1936: 1932: 1925: 1922: 1910: 1906: 1900: 1896: 1895: 1887: 1884: 1872: 1868: 1864: 1860: 1854: 1850: 1849: 1841: 1838: 1833: 1827: 1823: 1818: 1817: 1808: 1805: 1800: 1796: 1792: 1788: 1784: 1780: 1776: 1772: 1771: 1763: 1761: 1759: 1757: 1755: 1753: 1751: 1749: 1747: 1745: 1743: 1741: 1739: 1737: 1735: 1733: 1731: 1729: 1727: 1725: 1723: 1721: 1719: 1717: 1715: 1713: 1711: 1709: 1707: 1705: 1701: 1694: 1689: 1679: 1676: 1669: 1665: 1662: 1660: 1659: 1655: 1653: 1650: 1648: 1645: 1643: 1642:Ophthalmology 1640: 1638: 1635: 1633: 1630: 1628: 1625: 1623: 1620: 1618: 1615: 1613: 1610: 1608: 1605: 1603: 1600: 1598: 1595: 1592: 1589: 1586: 1583: 1582: 1578: 1569: 1564: 1557: 1552: 1547: 1545: 1543: 1538: 1534: 1530: 1526: 1520: 1518: 1514: 1510: 1503: 1501: 1499: 1494: 1490: 1486: 1485:colour vision 1481: 1479: 1475: 1471: 1467: 1463: 1458: 1456: 1452: 1448: 1440: 1438: 1434: 1431: 1427: 1423: 1418: 1414: 1413:Colour vision 1406: 1404: 1400: 1399:) in humans. 1398: 1397:normal vision 1394: 1388: 1386: 1381: 1376: 1374: 1370: 1369: 1363: 1359: 1358:Visual acuity 1353: 1350:The eye of a 1348: 1342:Visual acuity 1341: 1336: 1334: 1331: 1327: 1323: 1320: 1315: 1312: 1306: 1302: 1299: 1290: 1288: 1286: 1282: 1278: 1272: 1270: 1266: 1260: 1258: 1254: 1248: 1245: 1240: 1238: 1234: 1230: 1226: 1222: 1217: 1214: 1210: 1206: 1199: 1194: 1189: 1181: 1179: 1177: 1173: 1168: 1163: 1160: 1152: 1150: 1148: 1144: 1140: 1139: 1133: 1131: 1127: 1126: 1120: 1118: 1114: 1108: 1106: 1103:) absorb the 1102: 1098: 1094: 1089: 1085: 1081: 1077: 1072: 1070: 1066: 1062: 1054: 1052: 1046: 1044: 1042: 1038: 1034: 1030: 1026: 1022: 1018: 1009: 1006: 1003: 1002: 1001: 995: 993: 991: 985: 983: 982: 977: 973: 969: 961: 959: 955: 953: 952:phased arrays 949: 945: 941: 933: 928: 920: 914: 909: 904: 903:Arthropod eye 898: 891:Compound eyes 890: 888: 886: 883: 879: 874: 872: 871: 866: 862: 858: 853: 846: 844: 842: 838: 834: 826: 822: 818: 814: 810: 808: 807:Luneburg lens 804: 800: 792: 787: 780: 778: 774: 772: 771: 766: 765: 761: 753: 751: 748: 743: 739: 737: 734:, which have 733: 729: 725: 721: 717: 712: 710: 709:box jellyfish 705: 703: 698: 690: 688: 686: 682: 678: 674: 670: 666: 662: 660: 652: 650: 648: 644: 640: 636: 632: 624: 622: 620: 616: 611: 609: 605: 601: 597: 592: 590: 586: 582: 578: 574: 570: 566: 565:compound eyes 557: 555: 553: 549: 545: 541: 537: 533: 529: 525: 521: 517: 513: 509: 508:field of view 505: 500: 497: 493: 489: 487: 486:hyperspectral 483: 482:mantis shrimp 475: 474: 468: 464: 462: 457: 452: 450: 446: 442: 438: 434: 430: 426: 422: 418: 414: 410: 406: 402: 398: 394: 391:known as the 390: 386: 382: 378: 374: 368: 362: 360: 357:, which have 356: 352: 348: 344: 340: 336: 328: 324: 320: 314: 309: 302: 300: 298: 294: 290: 286: 282: 277: 273: 271: 267: 263: 262:compound eyes 258: 256: 255:visual cortex 252: 248: 244: 240: 236: 232: 228: 223: 221: 220:visual system 217: 213: 209: 205: 201: 200:sensory organ 197: 185: 179: 175: 172: 168: 165: 163: 159: 156: 152: 149: 147: 143: 140: 137: 135: 131: 126: 123: 120: 118: 114: 109: 105: 102:Diagram of a 99: 94: 90: 86: 81: 76: 71: 68: 64: 60: 56: 49: 45: 44:Eye (cyclone) 41: 37: 33: 19: 4639: 4533:Ciliary body 4373:Diencephalon 4372: 4357:Parasol cell 4341:Bipolar cell 4192:Pars plicata 4174:Ciliary body 4042: 3907:the original 3902: 3870:Plenum Press 3868:. New York: 3865: 3858:Bibliography 3812: 3808: 3754: 3750: 3696: 3690: 3664:. Retrieved 3639: 3635: 3622: 3597: 3589: 3577: 3566:. Retrieved 3549: 3545: 3535: 3524:. Retrieved 3496: 3489: 3478:. Retrieved 3469: 3460: 3449:. Retrieved 3421: 3414: 3406: 3400:. Retrieved 3372: 3365: 3360:, p. 28 3353: 3320: 3314: 3289: 3285: 3260: 3231:. Retrieved 3227:the original 3217: 3176: 3170: 3164: 3129: 3123: 3102: 3097: 3052: 3048: 3038: 3005: 3001: 2997: 2993: 2987: 2962: 2958: 2952: 2933: 2927: 2915: 2870: 2866: 2856: 2831: 2827: 2820: 2809:. Retrieved 2781: 2775: 2766: 2730: 2726: 2720: 2708:. Retrieved 2701:the original 2692: 2685: 2658: 2654: 2644: 2611: 2607: 2601: 2576: 2572: 2568: 2562: 2543: 2539: 2529: 2517:. Retrieved 2510:the original 2489: 2485: 2472: 2461:the original 2448: 2435: 2392: 2388: 2378: 2353: 2349: 2295: 2291: 2281: 2270: 2259: 2234: 2228: 2222: 2210:. Retrieved 2196: 2155: 2151: 2145: 2128: 2124: 2118: 2073: 2069: 2056: 2037: 2033: 1991: 1987: 1983: 1977: 1965:. Retrieved 1952: 1943: 1935:the original 1924: 1913:. Retrieved 1893: 1886: 1875:. Retrieved 1847: 1840: 1815: 1807: 1774: 1768: 1678: 1656: 1627:Eye movement 1532: 1524: 1521: 1511: 1507: 1504:Pigmentation 1482: 1459: 1444: 1435: 1419: 1416: 1401: 1389: 1377: 1365: 1356: 1332: 1328: 1324: 1307: 1303: 1294: 1273: 1261: 1249: 1241: 1221:bird of prey 1218: 1202: 1166: 1164: 1159:ciliary body 1158: 1156: 1143:brittle star 1141:, a type of 1136: 1135:The body of 1134: 1123: 1121: 1116: 1109: 1101:optical axes 1073: 1064: 1061:Strepsiptera 1058: 1050: 1013: 999: 986: 979: 965: 956: 944:polarisation 937: 897:Compound eye 875: 868: 854: 850: 830: 811: 796: 775: 768: 762: 757: 744: 740: 713: 706: 694: 685:ion channels 683:but TRP are 663: 656: 628: 612: 593: 561: 538:. They have 520:caterpillars 501: 490: 479: 471: 461:Strepsiptera 453: 363: 332: 278: 274: 259: 224: 206:to perceive 195: 193: 155:A01.1.00.007 151:A15.2.00.001 138: 85:Compound eye 67: 18:Eye membrane 4625:Eye disease 4605:Keratocytes 4494:Conjunctiva 4451:Ora serrata 4389:Muller glia 4353:Midget cell 4073:Endothelium 4063:Dua's layer 3930:Adobe Flash 2922:, p. 8 2710:13 November 2492:: 147–177. 2264:Animal Eyes 1617:Eye disease 1498:optic nerve 1366:cycles per 1319:Glyptonotus 1229:ultraviolet 1198:mollusc eye 1093:pseudopupil 1080:dragonflies 1004:refracting, 958:ommatidia. 948:diffraction 728:cephalopods 643:crustaceans 635:cephalopods 631:vertebrates 604:entrainment 577:rhabdomeric 433:cephalopods 413:transparent 409:optic nerve 381:vertebrates 270:crustaceans 251:optic nerve 241:to form an 128:Identifiers 4660:Categories 4466:of the eye 4439:Optic disc 4197:Pars plana 4068:Descemet's 4048:Epithelium 3666:2008-09-16 3598:The Senses 3568:2012-01-01 3546:J Exp Biol 3526:2020-10-19 3480:2015-06-03 3451:2020-10-19 3402:2020-10-19 3233:2012-09-01 2998:Drosophila 2811:2008-09-15 2579:(2): 137. 2573:Alcheringa 2230:BioScience 1915:2020-10-19 1877:2020-10-19 1848:Psychology 1690:References 1622:Eye injury 1587:(focusing) 1540:r-opsins. 1478:blind spot 1426:cone cells 1362:cone cells 1337:Physiology 1269:crystallin 1069:trilobites 990:ommatidium 940:ommatidium 716:gastropods 665:Pit vipers 585:gastropods 492:Trilobites 456:arthropods 441:amphibians 397:cone cells 379:. 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