Eye - Wikipedia PDF

3/26/2019 Eye - Wikipedia
Eye
Ey es are organs of the v isual sy stem. They prov ide organisms with
Eye
v ision, the ability to receiv e and process v isual detail, as well as
enabling sev eral photo response functions that are independent of
v ision. Ey es detect light and conv ert it into electro-chemical impulses
in neurons. In higher organisms, the ey e is a complex optical sy stem
which collects light from the surrounding env ironment, regulates its
intensity through a diaphragm, focuses it through an adjustable
assembly of lenses to form an image, conv erts this image into a set of
electrical signals, and transmits these signals to the brain through
complex neural pathway s that connect the ey e v ia the optic nerv e to
the v isual cortex and other areas of the brain. Ey es with resolv ing
power hav e come in ten fundamentally different forms, and 96% of
animal species possess a complex optical sy stem. [1 ] Image-resolv ing
ey es are present in molluscs, chordates and arthropods. [2 ]
Human eye
The simplest "ey es", such as those in microorganisms, do nothing but
detect whether the surroundings are light or dark, which is sufficient for
the entrainment of circadian rhy thms. [3 ] From more complex ey es,
retinal photosensitiv e ganglion cells send signals along the
retinohy pothalamic tract to the suprachiasmatic nuclei to effect
circadian adjustment and to the pretectal area to control the pupillary
light reflex.
Contents
Overview
Types
Compound eye of Antarctic krill
Non-compound eyes Details
Pit eyes
System Nervous
Spherical lens eye
Multiple lenses Identifiers
Refractive cornea Latin oculus
Reflector eyes
MeSH D005123 (https://meshb.nlm.n
Compound eyes ih.gov/record/ui?ui=D005123)
Apposition eyes
Superposition eyes TA A15.2.00.001 (http://www.unifr.
Parabolic superposition ch/ifaa/Public/EntryPage/TA9
Other 8%20Tree/Entity%20TA98%2
Nutrients 0EN/15.2.00.001%20Entity%2
0TA98%20EN.htm)
Evolution
Relationship to life requirements
A01.1.00.007 (http://www.unifr.
https://en.wikipedia.org/wiki/Eye 1/16
Physiology ch/ifaa/Public/EntryPage/TA9
Visual acuity 8%20Tree/Entity%20TA98%2
Colour perception 0EN/01.1.00.007%20Entity%2
Rods and cones 0TA98%20EN.htm)
Pigmentation FMA 54448 75665, 54448 (https://
Additional images bioportal.bioontology.org/ontol
See also ogies/FMA/?p=classes&conce
Notes ptid=http%3A%2F%2Fpurl.or
References g%2Fsig%2Font%2Ffma%2Ff
Bibliography ma75665,)
Further reading Anatomical terminology
External links
Overview
Complex ey es can distinguish shapes and
colours. The v isual fields of many
organisms, especially predators, inv olv e
large areas of binocular v ision to improv e
depth perception. In other organisms,
ey es are located so as to maximise the
field of v iew, such as in rabbits and
horses, which hav e monocular v ision.
Eye of European bison Human eye
The first proto-ey es ev olv ed among
animals 600 million y ears ago about the
time of the Cambrian explosion. [4 ] The last common ancestor of animals possessed the biochemical toolkit
necessary for v ision, and more adv anced ey es hav e ev olv ed in 96% of animal species in six of the ~35 [a ] main
phy la. [1 ] In most v ertebrates and some molluscs, the ey e works by allowing light to enter and project onto a light-
sensitiv e panel of cells, known as the retina, at the rear of the ey e. The cone cells (for colour) and the rod cells (for
low-light contrasts) in the retina detect and conv ert light into neural signals for v ision. The v isual signals are then
transmitted to the brain v ia the optic nerv e. Such ey es are ty pically roughly spherical, filled with a transparent gel-
like substance called the v itreous humour, with a focusing lens and often an iris; the relaxing or tightening of the
muscles around the iris change the size of the pupil, thereby regulating the amount of light that enters the ey e, [5 ]
and reducing aberrations when there is enough light. [6 ] The ey es of most cephalopods, fish, amphibians and snakes
hav e fixed lens shapes, and focusing v ision is achiev ed by telescoping the lens—similar to how a camera focuses. [7 ]
Compound ey es are found among the arthropods and are composed of many simple facets which, depending on the
details of anatomy , may giv e either a single pixelated image or multiple images, per ey e. Each sensor has its own lens
and photosensitiv e cell(s). Some ey es hav e up to 28,000 such sensors, which are arranged hexagonally , and which
can giv e a full 360° field of v ision. Compound ey es are v ery sensitiv e to motion. Some arthropods, including many
Strepsiptera, hav e compound ey es of only a few facets, each with a retina capable of creating an image, creating
v ision. With each ey e v iewing a different thing, a fused image from all the ey es is produced in the brain, prov iding
v ery different, high-resolution images.
Possessing detailed hy perspectral colour v ision, the Mantis shrimp has been reported to hav e the world's most
complex colour v ision sy stem. [8 ] Trilobites, which are now extinct, had unique compound ey es. They used clear
calcite cry stals to form the lenses of their ey es. In this, they differ from most other arthropods, which hav e soft ey es.
The number of lenses in such an ey e v aried, howev er: some trilobites had only one, and some had thousands of
lenses in one ey e.
In contrast to compound ey es, simple ey es are those that hav e a single lens. For example, jumping spiders hav e a
large pair of simple ey es with a narrow field of v iew, supported by an array of other, smaller ey es for peripheral
v ision. Some insect larv ae, like caterpillars, hav e a different ty pe of simple ey e (stemmata) which usually prov ides
only a rough image, but (as in sawfly larv ae) can possess resolv ing powers of 4 degrees of arc, be polarization
sensitiv e and capable of increasing its absolute sensitiv ity at night by a factor of 1,000 or more. [9 ] Some of the
simplest ey es, called ocelli, can be found in animals like some of the snails, which cannot actually "see" in the normal
sense. They do hav e photosensitiv e cells, but no lens and no other means of projecting an image onto these cells.
They can distinguish between light and dark, but no more. This enables snails to keep out of direct sunlight. In
organisms dwelling near deep-sea v ents, compound ey es hav e been secondarily simplified and adapted to spot the
infra-red light produced by the hot v ents—in this way the bearers can spot hot springs and av oid being boiled
aliv e. [1 0 ]
Types
There are ten different ey e lay outs—indeed ev ery technological method of capturing an optical image commonly
used by human beings, with the exceptions of zoom and Fresnel lenses, occur in nature. [1 ] Ey e ty pes can be
categorised into "simple ey es", with one concav e photoreceptiv e surface, and "compound ey es", which comprise a
number of indiv idual lenses laid out on a conv ex surface. [1 ] Note that "simple" does not imply a reduced lev el of
complexity or acuity . Indeed, any ey e ty pe can be adapted for almost any behav iour or env ironment. The only
limitations specific to ey e ty pes are that of resolution—the phy sics of compound ey es prev ents them from achiev ing
a resolution better than 1°. Also, superposition ey es can achiev e greater sensitiv ity than apposition ey es, so are
better suited to dark-dwelling creatures. [1 ] Ey es also fall into two groups on the basis of their photoreceptor's
cellular construction, with the photoreceptor cells either being cilliated (as in the v ertebrates) or rhabdomeric.
These two groups are not monophy letic; the cnidaria also possess cilliated cells, [1 1 ] and some gastropods, [1 2 ] as
well as some annelids possess both. [1 3 ]
Non-compound eyes
Simple ey es are rather ubiquitous, and lens-bearing ey es hav e ev olv ed at least sev en times in v ertebrates,
cephalopods, annelids, crustaceans and cubozoa. [1 4 ]
Pit eyes
Pit ey es, also known as stemma, are ey e-spots which may be set into a pit to reduce the angles of light that enters and
affects the ey e-spot, to allow the organism to deduce the angle of incoming light. [1 ] Found in about 85% of phy la,
these basic forms were probably the precursors to more adv anced ty pes of "simple ey es". They are small, comprising
up to about 100 cells cov ering about 100 µm. [1 ] The directionality can be improv ed by reducing the size of the
aperture, by incorporating a reflectiv e lay er behind the receptor cells, or by filling the pit with a refractile
material. [1 ]
Pit v ipers hav e dev eloped pits that function as ey es by sensing thermal infra-red radiation, in addition to their
optical wav elength ey es like those of other v ertebrates.
Spherical lens eye

The resolution of pit ey es can be greatly improv ed by incorporating a material with a higher refractiv e index to form
a lens, which may greatly reduce the blur radius encountered—hence increasing the resolution obtainable. [1 ] The
most basic form, seen in some gastropods and annelids, consists of a lens of one refractiv e index. A far sharper image
can be obtained using materials with a high refractiv e index, decreasing to the edges; this decreases the focal length
and thus allows a sharp image to form on the retina. [1 ] This also allows a larger aperture for a giv en sharpness of
image, allowing more light to enter the lens; and a flatter lens, reducing spherical aberration. [1 ] Such a non-
homogeneous lens is necessary for the focal length to drop from about 4 times the lens radius, to 2.5 radii. [1 ]
Heterogeneous ey es hav e ev olv ed at least nine times: four or more times in gastropods, once in the copepods, once
in the annelids, once in the cephalopods, [1 ] and once in the chitons, which hav e aragonite lenses. [1 5 ] No extant
aquatic organisms possess homogeneous lenses; presumably the ev olutionary pressure for a heterogeneous lens is
great enough for this stage to be quickly "outgrown". [1 ]
This ey e creates an image that is sharp enough that motion of the ey e can cause significant blurring. To minimise the
effect of ey e motion while the animal mov es, most such ey es hav e stabilising ey e muscles. [1 ]
The ocelli of insects bear a simple lens, but their focal point alway s lies behind the retina; consequently they can
nev er form a sharp image. Ocelli (pit-ty pe ey es of arthropods) blur the image across the whole retina, and are
consequently excellent at responding to rapid changes in light intensity across the whole v isual field; this fast
response is further accelerated by the large nerv e bundles which rush the information to the brain. [1 6 ] 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 sensitiv ity . [1 6 ] 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 v egetation, usually
comes from abov e). [1 6 ]
Multiple lenses
Some marine organisms bear more than one lens; for instance the copepod Pontella has three. The outer has a
parabolic surface, countering the effects of spherical aberration while allowing a sharp image to be formed. Another
copepod, Copilia, has two lenses in each ey e, arranged like those in a telescope. [1 ] Such arrangements are rare and
poorly understood, but represent an alternativ e construction.
Multiple lenses are seen in some hunters such as eagles and jumping spiders, which hav e a refractiv e cornea: these
hav e a negativ e lens, enlarging the observ ed image by up to 50% ov er the receptor cells, thus increasing their optical
resolution. [1 ]
Refractive cornea
In the ey es of most mammals, birds, reptiles, and most other terrestrial v ertebrates (along with spiders and some
insect larv ae) the v itreous fluid has a higher refractiv e index than the air. [1 ] In general, the lens is not spherical.
Spherical lenses produce spherical aberration. In refractiv e corneas, the lens tissue is corrected with
inhomogeneous lens material (see Luneburg lens), or with an aspheric shape. [1 ] Flattening the lens has a
disadv antage; the quality of v ision is diminished away from the main line of focus. Thus, animals that hav e ev olv ed
with a wide field-of-v iew often hav e ey es that make use of an inhomogeneous lens. [1 ]
As mentioned abov e, a refractiv e cornea is only useful out of water. In water, there is little difference in refractiv e
index between the v itreous fluid and the surrounding water. Hence creatures that hav e returned to the water—
penguins and seals, for example—lose their highly curv ed cornea and return to lens-based v ision. An alternativ e
solution, borne by some div ers, is to hav e a v ery strongly focusing cornea. [1 ]
Reflector eyes
An alternativ e to a lens is to line the inside of the ey e with "mirrors", and reflect the image to focus at a central
point. [1 ] The nature of these ey es means that if one were to peer into the pupil of an ey e, one would see the same
image that the organism would see, reflected back out. [1 ]
Many small organisms such as rotifers, copepods and flatworms use such organs, but these are too small to produce
usable images. [1 ] Some larger organisms, such as scallops, also use reflector ey es. The scallop Pecten has up to 100
millimetre-scale reflector ey es fringing the edge of its shell. It detects mov ing objects as they pass successiv e
lenses. [1 ]
There is at least one v ertebrate, the spookfish, whose ey es include reflectiv e optics for focusing of light. Each of the
two ey es of a spookfish collects light from both abov e and below; the light coming from abov e is focused by a lens,
while that coming from below, by a curv ed mirror composed of many lay ers of small reflectiv e plates made of
guanine cry stals. [1 7 ]
Compound eyes
A compound ey e may consist of thousands of indiv idual photoreceptor
units or ommatidia (ommatidium, singular). The image perceiv ed is a
combination of inputs from the numerous ommatidia (indiv idual "ey e
units"), which are located on a conv ex surface, thus pointing in slightly
different directions. Compared with simple ey es, compound ey es possess a
v ery large v iew angle, and can detect fast mov ement and, in some cases,
the polarisation of light. [1 8 ] Because the indiv idual lenses are so small, the
effects of diffraction impose a limit on the possible resolution that can be
obtained (assuming that they do not function as phased array s). This can
An image of a house fly compound
only be countered by increasing lens size and number. To see with a eye surface by using scanning
resolution comparable to our simple ey es, humans would require v ery electron microscope
large compound ey es, around 11 metres (36 ft) in radius. [1 9 ]
Compound ey es fall into two groups: apposition ey es, which form multiple
inv erted images, and superposition ey es, which form a single erect image. [2 0 ]
Compound ey es are common in arthropods, and are also present in annelids
and some biv alv ed molluscs. [2 1 ] Compound ey es in arthropods grow at their
margins by the addition of new ommatidia. [2 2 ]
Apposition eyes
Apposition ey es are the most common form of ey es, and are presumably the
Anatomy of the compound eye of
ancestral form of compound ey es. They are found in all arthropod groups,
an insect
although they may hav e ev olv ed more than once within this phy lum. [1 ] Some
annelids and biv alv es also hav e apposition ey es. They are also possessed by
Limulus, the horseshoe crab, and there are suggestions that other
chelicerates dev eloped their simple ey es by reduction from a compound
starting point. [1 ] (Some caterpillars appear to hav e ev olv ed compound
ey es from simple ey es in the opposite fashion.)
Apposition ey es work by gathering a number of images, one from each ey e,

and combining them in the brain, with each ey e ty pically contributing a
single point of information. The ty pical apposition ey e has a lens focusing
light from one direction on the rhabdom, while light from other directions
is absorbed by the dark wall of the ommatidium. Arthropods such as this Calliphora
vomitoria fly have compound eyes
Superposition eyes
The second ty pe is named the superposition ey e. The superposition ey e is div ided into three ty pes:
refracting,
reflecting and
parabolic superposition
The refracting superposition ey e 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
ey e, which is where the tips of the rhabdoms are. This ty pe of compound ey e, for which a minimal size exists below
which effectiv e superposition cannot occur, [2 3 ] is normally found in nocturnal insects, because it can create images
up to 1000 times brighter than equiv alent apposition ey es, though at the cost of reduced resolution. [2 4 ] In the
parabolic superposition compound ey e ty pe, seen in arthropods such as may flies, the parabolic surfaces of the
inside of each facet focus light from a reflector to a sensor array . Long-bodied decapod crustaceans such as shrimp,
prawns, cray fish and lobsters are alone in hav ing reflecting superposition ey es, which also hav e a transparent gap
but use corner mirrors instead of lenses.
Parabolic superposition
This ey e ty pe functions by refracting light, then using a parabolic mirror to focus the image; it combines features of
superposition and apposition ey es. [1 0 ]
Other
Another kind of compound ey e, found in males of Order Strepsiptera, employ s a series of simple ey es—ey es hav ing
one opening that prov ides light for an entire image-forming retina. Sev eral of these eyelets together form the
strepsipteran compound ey e, which is similar to the 'schizochroal' compound ey es of some trilobites. [2 5 ] Because
each ey elet is a simple ey e, it produces an inv erted image; those images are combined in the brain to form one
unified image. Because the aperture of an ey elet is larger than the facets of a compound ey e, this arrangement allows
v ision under low light lev els. [1 ]
Good fliers such as flies or honey bees, or prey -catching insects such as pray ing mantis or dragonflies, hav e
specialised zones of ommatidia organised into a fov ea area which giv es acute v ision. In the acute zone, the ey es are
flattened and the facets larger. The flattening allows more ommatidia to receiv e light from a spot and therefore
higher resolution. The black spot that can be seen on the compound ey es of such insects, which alway s seems to look
directly at the observ er, is called a pseudopupil. This occurs because the ommatidia which one observ es "head-on"
(along their optical axes) absorb the incident light, while those to one side reflect it. [2 6 ]
There are some exceptions from the ty pes mentioned abov e. Some insects hav e a so-called single lens compound
ey e, a transitional ty pe which is something between a superposition ty pe of the multi-lens compound ey e and the
single lens ey e found in animals with simple ey es. Then there is the my sid shrimp, Dioptromysis paucispinosa. The
shrimp has an ey e of the refracting superposition ty pe, in the rear behind this in each ey e there is a single large facet
that is three times in diameter the others in the ey e and behind this is an enlarged cry stalline cone. This projects an
upright image on a specialised retina. The resulting ey e is a mixture of a simple ey e within a compound ey e.
Another v ersion is a compound ey e often referred to as "pseudofaceted", as seen in Scutigera. [2 7 ] This ty pe of ey e

consists of a cluster of numerous ommatidia on each side of the head, organised in a way that resembles a true
compound ey e.
The body of Ophiocoma wendtii, a ty pe of brittle star, is cov ered with ommatidia, turning its whole skin into a
compound ey e. The same is true of many chitons. The tube feet of sea urchins contain photoreceptor proteins, which
together act as a compound ey e; they lack screening pigments, but can detect the directionality of light by the
shadow cast by its opaque body . [2 8 ]
Nutrients
The ciliary body is triangular in horizontal section and is coated by a double lay er, the ciliary epithelium. The inner
lay er is transparent and cov ers the v itreous body , and is continuous from the neural tissue of the retina. The outer
lay er is highly pigmented, continuous with the retinal pigment epithelium, and constitutes the cells of the dilator
muscle.
The v itreous is the transparent, colourless, gelatinous mass that fills the space between the lens of the ey e and the
retina lining the back of the ey e. [2 9 ] It is produced by certain retinal cells. It is of rather similar composition to the
cornea, but contains v ery few cells (mostly phagocy tes which remov e unwanted cellular debris in the v isual field, as
well as the hy alocy tes of Balazs of the surface of the v itreous, which reprocess the hy aluronic acid), no blood v essels,
and 98–99% of its v olume is water (as opposed to 7 5% in the cornea) with salts, sugars, v itrosin (a ty pe of collagen),
a network of collagen ty pe II fibres with the mucopoly saccharide hy aluronic acid, and also a wide array of proteins
in micro amounts. Amazingly , with so little solid matter, it tautly holds the ey e.
Evolution
Photoreception is phy logenetically v ery old, with v arious theories of phy logenesis. [3 0 ] The common origin
(monophy ly ) of all animal ey es is now widely accepted as fact. This is based upon the shared genetic features of all
ey es; that is, all modern ey es, v aried as they are, hav e their origins in a proto-ey e believ ed to hav e ev olv ed some
540 million y ears ago, [3 1 ][3 2 ][3 3 ] and the PAX6 gene is considered a key factor in this. The majority of the
adv ancements in early ey es are believ ed to hav e taken only a few million y ears to dev elop, since the first predator to
gain true imaging would hav e touched off an "arms race"[3 4 ] among all species that did not flee the photopic
env ironment. Prey animals and competing predators alike would be at a distinct disadv antage without such
capabilities and would be less likely to surv iv e and reproduce. Hence multiple ey e ty pes and subty pes dev eloped in
parallel (except those of groups, such as the v ertebrates, that were only forced into the photopic env ironment at a
late stage).
Ey es in v arious animals show adaptation to their requirements. For example, the ey e of a bird of prey has much
greater v isual acuity than a human ey e, and in some cases can detect ultrav iolet radiation. The different forms of ey e
in, for example, v ertebrates and molluscs are examples of parallel ev olution, despite their distant common ancestry .
Phenoty pic conv ergence of the geometry of cephalopod and most v ertebrate ey es creates the impression that the
v ertebrate ey e ev olv ed from an imaging cephalopod ey e,

but this is not the case, as the rev ersed roles of their
respectiv e ciliary and rhabdomeric opsin classes[3 5 ] and
different lens cry stallins show. [3 6 ]
The v ery earliest "ey es", called ey e-spots, were simple

patches of photoreceptor protein in unicellular animals.
In multicellular beings, multicellular ey espots ev olv ed,
phy sically similar to the receptor patches for taste and
smell. These ey espots could only sense ambient
brightness: they could distinguish light and dark, but not
the direction of the light source. [1 ]
Through gradual change, the ey e-spots of species liv ing

in well-lit env ironments depressed into a shallow "cup"
shape, the ability to slightly discriminate directional
brightness was achiev ed by using the angle at which the
light hit certain cells to identify the source. The pit
deepened ov er time, the opening diminished in size, and
the number of photoreceptor cells increased, forming an
effectiv e pinhole camera that was capable of dimly
distinguishing shapes. [3 7 ] Howev er, the ancestors of
modern hagfish, thought to be the protov ertebrate [3 5 ]
were ev idently pushed to v ery deep, dark waters, where
Evolution of the mollusc eye
they were less v ulnerable to sighted predators, and
where it is adv antageous to hav e a conv ex ey e-spot,
which gathers more light than a flat or concav e one. This would hav e led to a somewhat different ev olutionary
trajectory for the v ertebrate ey e than for other animal ey es.
The thin ov ergrowth of transparent cells ov er the ey e's aperture, originally formed to prev ent damage to the
ey espot, allowed the segregated contents of the ey e chamber to specialise into a transparent humour that optimised
colour filtering, blocked harmful radiation, improv ed the ey e's refractiv e index, and allowed functionality outside of
water. The transparent protectiv e cells ev entually split into two lay ers, with circulatory fluid in between that
allowed wider v iewing angles and greater imaging resolution, and the thickness of the transparent lay er gradually
increased, in most species with the transparent cry stallin protein. [3 8 ]
The gap between tissue lay ers naturally formed a bioconv ex shape, an optimally ideal structure for a normal
refractiv e index. Independently , a transparent lay er and a nontransparent lay er split forward from the lens: the
cornea and iris. Separation of the forward lay er again formed a humour, the aqueous humour. This increased
refractiv e power and again eased circulatory problems. Formation of a nontransparent ring allowed more blood
v essels, more circulation, and larger ey e sizes. [3 8 ]
Relationship to life requirements

Ey es are generally adapted to the env ironment 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 liv e on the African plains, hav ing a horizontal line of high-density
ganglia, while tree-dwelling creatures which require good all-round v ision tend to hav e a sy mmetrical distribution of
ganglia, with acuity decreasing outwards from the centre.
Of course, for most ey e ty pes, it is impossible to div erge from a spherical form, so only the density of optical
receptors can be altered. In organisms with compound ey es, it is the number of ommatidia rather than ganglia that
reflects the region of highest data acquisition. [1 ]:2 3 –2 4 Optical superposition ey es are constrained to a spherical
shape, but other forms of compound ey es may deform to a shape where more ommatidia are aligned to, say , the
horizon, without altering the size or density of indiv idual ommatidia. [3 9 ] Ey es of horizon-scanning organisms hav e
stalks so they can be easily aligned to the horizon when this is inclined, for example if the animal is on a slope. [2 6 ]
An extension of this concept is that the ey es of predators ty pically hav e a zone of v ery acute v ision at their centre, to
assist in the identification of prey . [3 9 ] In deep water organisms, it may not be the centre of the ey e that is enlarged.
The hy periid amphipods are deep water animals that feed on organisms abov e them. Their ey es are almost div ided
into two, with the upper region thought to be inv olv ed in detecting the silhouettes of potential prey —or predators—
against the faint light of the sky abov e. Accordingly , deeper water hy periids, where the light against which the
silhouettes must be compared is dimmer, hav e larger "upper-ey es", and may lose the lower portion of their ey es
altogether. [3 9 ] In the giant Antarctic isopod Gly ptonotus a small v entral compound ey e is phy sically completely
separated from the much larger dorsal compound ey e. [4 0 ] Depth perception can be enhanced by hav ing ey es which
are enlarged in one direction; distorting the ey e slightly allows the distance to the object to be estimated with a high
degree of accuracy . [1 0 ]
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 v ery large backdrop. [3 9 ] On the other hand, the ey es of organisms which operate in low light lev els,
such as around dawn and dusk or in deep water, tend to be larger to increase the amount of light that can be
captured. [3 9 ]
It is not only the shape of the ey e that may be affected by lifesty le. Ey es can be the most v isible parts of organisms,
and this can act as a pressure on organisms to hav e more transparent ey es at the cost of function. [3 9 ]
Ey es may be mounted on stalks to prov ide better all-round v ision, by lifting them abov e an organism's carapace; this
also allows them to track predators or prey without mov ing the head. [1 0 ]
Physiology
Visual acuity
Visual acuity , or resolv ing power, is "the ability to distinguish fine detail" and is the property of cone cells. [4 1 ] It is
often measured in cycles per degree (CPD), which measures an angular resolution, or how much an ey e can
differentiate one object from another in terms of v isual angles. Resolution in CPD can be measured by bar charts of
different numbers of white/black stripe cy cles. For example, if each pattern is 1.7 5 cm wide and is placed at 1 m
distance from the ey e, 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 cy cles per degree of that pattern. The highest such number that the ey e can resolv e as stripes, or
distinguish from a grey block, is then the measurement of v isual acuity of the ey e.
For a human ey e with excellent acuity , the maximum theoretical resolution is 50 CPD[4 2 ] (1.2 arcminute per line
pair, or a 0.35 mm line pair, at 1 m). A rat can resolv e only about 1 to 2 CPD. [4 3 ] A horse has higher acuity through
most of the v isual field of its ey es than a human has, but does not match the high acuity of the human ey e's central
fov ea region. [4 4 ]
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 improv ed resolution of
approximately 1.7 arcminutes per line pair. [4 5 ] A resolution of 2
arcminutes per line pair, equiv alent to a 1 arcminute gap in an optoty pe,
corresponds to 20/20 (normal v ision) in humans.
Howev er, in the compound ey e, the resolution is related to the size of

indiv idual ommatidia and the distance between neighbouring ommatidia.
Phy sically these cannot be reduced in size to achiev e the acuity seen with
single lensed ey es as in mammals. Compound ey es hav e a much lower
acuity than v ertebrate ey es. [4 6 ]
The eye of a red-tailed hawk
Colour perception
"Colour v ision is the faculty of the organism to distinguish lights of different spectral qualities."[4 7 ] All organisms are
restricted to a small range of electromagnetic spectrum; this v aries from creature to creature, but is mainly between
wav elengths of 400 and 7 00 nm. [4 8 ] This is a rather small section of the electromagnetic spectrum, probably
reflecting the submarine ev olution of the organ: water blocks out all but two small windows of the EM spectrum, and
there has been no ev olutionary pressure among land animals to broaden this range. [4 9 ]
The most sensitiv e pigment, rhodopsin, has a peak response at 500 nm. [5 0 ] Small changes to the genes coding for
this protein can tweak the peak response by a few nm;[2 ] pigments in the lens can also filter incoming light, changing
the peak response. [2 ] Many organisms are unable to discriminate between colours, seeing instead in shades of grey ;
colour v ision necessitates a range of pigment cells which are primarily sensitiv e to smaller ranges of the spectrum.
In primates, geckos, and other organisms, these take the form of cone cells, from which the more sensitiv e rod cells
ev olv ed. [5 0 ] Ev en if organisms are phy sically capable of discriminating different colours, this does not necessarily
mean that they can perceiv e the different colours; only with behav ioural tests can this be deduced. [2 ]
Most organisms with colour v ision are able to detect ultrav iolet light. This high energy light can be damaging to
receptor cells. With a few exceptions (snakes, placental mammals), most organisms av oid these effects by hav ing
absorbent oil droplets around their cone cells. The alternativ e, dev eloped by organisms that had lost these oil
droplets in the course of ev olution, is to make the lens imperv ious to UV light—this precludes the possibility of any
UV light being detected, as it does not ev en reach the retina. [5 0 ]
Rods and cones

The retina contains two major ty pes of light-sensitiv e photoreceptor cells used for v ision: the rods and the cones.
Rods cannot distinguish colours, but are responsible for low-light (scotopic) monochrome (black-and-white) v ision;
they work well in dim light as they contain a pigment, rhodopsin (v isual purple), which is sensitiv e at low light
intensity , but saturates at higher (photopic) intensities. Rods are distributed throughout the retina but there are
none at the fov ea and none at the blind spot. Rod density is greater in the peripheral retina than in the central retina.
Cones are responsible for colour v ision. They require brighter light to function than rods require. In humans, there
are three ty pes of cones, maximally sensitiv e to long-wav elength, medium-wav elength, and short-wav elength light
(often referred to as red, green, and blue, respectiv ely , though the sensitiv ity peaks are not actually at these
colours). The colour seen is the combined effect of stimuli to, and responses from, these three ty pes of cone cells.
Cones are mostly concentrated in and near the fov ea. Only a few are present at the sides of the retina. Objects are
seen most sharply in focus when their images fall on the fov ea, as when one looks at an object directly . Cone cells and
rods are connected through intermediate cells in the retina to nerv e fibres of the optic nerv e. When rods and cones
are stimulated by light, they connect through adjoining cells within the retina to send an electrical signal to the optic
nerv e fibres. The optic nerv es send off impulses through these fibres to the brain. [5 0 ]
Pigmentation
The pigment molecules used in the ey e are v arious, but can be used to define the ev olutionary distance between
different groups, and can also be an aid in determining which are closely related—although problems of conv ergence
do exist. [5 0 ]
Opsins are the pigments inv olv ed in photoreception. Other pigments, such as melanin, are used to shield the
photoreceptor cells from light leaking in from the sides. The opsin protein group ev olv ed long before the last
common ancestor of animals, and has continued to div ersify since. [2 ]
There are two ty pes of opsin inv olv ed in v ision; c-opsins, which are associated with ciliary -ty pe photoreceptor cells,
and r-opsins, associated with rhabdomeric photoreceptor cells. [5 1 ] The ey es of v ertebrates usually contain cilliary
cells with c-opsins, and (bilaterian) inv ertebrates hav e rhabdomeric cells in the ey e with r-opsins. Howev er, some
ganglion cells of v ertebrates express r-opsins, suggesting that their ancestors used this pigment in v ision, and that
remnants surv iv e in the ey es. [5 1 ] Likewise, c-opsins hav e been found to be expressed in the brain of some
inv ertebrates. They may hav e been expressed in ciliary cells of larv al ey es, which were subsequently resorbed into
the brain on metamorphosis to the adult form. [5 1 ] C-opsins are also found in some deriv ed bilaterian-inv ertebrate
ey es, such as the pallial ey es of the biv alv e molluscs; howev er, the lateral ey es (which were presumably the ancestral
ty pe for this group, if ey es ev olv ed once there) alway s use r-opsins. [5 1 ] Cnidaria, which are an outgroup to the taxa
mentioned abov e, express c-opsins—but r-opsins are y et to be found in this group. [5 1 ] Incidentally , the melanin
produced in the cnidaria is produced in the same fashion as that in v ertebrates, suggesting the common descent of
this pigment. [5 1 ]
Additional images
The structures of the eye Another view of the eye

labelled and the structures of the
eye labelled
See also
Adaptation (eye) (Night vision)
Emission theory (vision)

Eye colour
Eye development
Eye disease
Eye injury
Eye movement
Eyelid
Nictitating membrane
Orbital x-ray
Simple eye in invertebrates
Tapetum lucidum
Tears
Notes
a. 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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Bibliography
Ali, Mohamed Ather; Klyne, M.A. (1985). Vision in Vertebrates. New York: Plenum Press. ISBN 978-0-306-42065-8.
Further reading
Yong, Ed (14 January 2016). "Inside the Eye: Nature's Most Exquisite Creation" (http://ngm.nationalgeographic.com/201
6/02/evolution-of-eyes-text). National Geographic.
External links
Evolution of the eye (https://www.pbs.org/wgbh/evolution/library/01/1/l_011_01.html)
Anatomy of the eye – flash animated interactive. (http://www.lensshopper.com/eye-anatomy.asp) (Adobe Flash)
Webvision. The organisation of the retina and visual system. (http://webvision.med.utah.edu/) An in-depth treatment of
retinal function, open to all but geared most towards graduate students.
Eye strips images of all but bare essentials before sending visual information to brain, UC Berkeley research shows (htt
p://www.berkeley.edu/news/media/releases/2001/03/28_wers1.html)
Retrieved from "https://en.wikipedia.org/w/index.php?title=Eye&oldid=886983163"
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Spherical lens eye

Apposition ey es work by gathering a number of images, one from each ey e,

Another v ersion is a compound ey e often referred to as "pseudofaceted", as seen in Scutigera. [2 7 ] This ty pe of ey e

v ertebrate ey e ev olv ed from an imaging cephalopod ey e,

The v ery earliest "ey es", called ey e-spots, were simple

Through gradual change, the ey e-spots of species liv ing

Relationship to life requirements

Spherical aberration limits the resolution of a 7 mm pupil to about 3

Howev er, in the compound ey e, the resolution is related to the size of

Rods and cones

The structures of the eye Another view of the eye

Emission theory (vision)

41. Ali & Klyne 1985, p. 28

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This page was last edited on 9 March 2019, at 20:59 (UTC).

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