The pupil is a hole located in the centre of the iris of the eye that allows light to strike theretina.

[1] It
appears black because light rays entering the pupil are either absorbed by thetissues inside the eye
directly, or absorbed after diffuse reflections within the eye that mostly miss exiting the narrow pupil.
In humans the pupil is round, but other species, such as some cats, have vertical slit
pupils,goats have horizontally oriented pupils, and some catfish have annular types.[2] In optical
terms, the anatomical pupil is the eye's aperture and the iris is the aperture stop. The image of the
pupil as seen from outside the eye is the entrance pupil, which does not exactly correspond to the
location and size of the physical pupil because it is magnified by thecornea. On the inner edge lies a
prominent structure, the collarette, marking the junction of the embryonic pupillary membrane
covering the embryonic pupil.

Controlling[edit]
The iris is a contractile structure, consisting mainly of smooth muscle, surrounding the pupil. Light
enters the eye through the pupil, and the iris regulates the amount of light by controlling the size of
the pupil. The iris contains two groups of smooth muscles; a circular group called the sphincter
pupillae, and a radial group called the dilator pupillae. When the sphincter pupillae contract, the iris
decreases or constricts the size of the pupil. The dilator pupillae, innervated by sympathetic nerves
from the superior cervical ganglion, cause the pupil to dilate when they contract. These muscles are
sometimes referred to as intrinsic eye muscles. The sensory pathway (rod or cone, bipolar, ganglion)
is linked with its counterpart in the other eye by a partial crossover of each eye's fibers. This causes
the effect in one eye to carry over to the other. If the drug pilocarpine is administered, the pupils will
constrict and accommodation is increased due to the parasympathetic action on the circular muscle
fibers, conversely, atropine will cause paralysis of accommodation (cycloplegia) and dilation of the
pupil.

Optic effects[edit]
When bright light is shone on the eye, light sensitive cells in the retina, including rod and cone
photoreceptors and melanopsin ganglion cells, will send signals to the oculomotor nerve, specifically
the parasympathetic part coming from the Edinger-Westphal nucleus, which terminates on the
circular iris sphincter muscle. When this muscle contracts, it reduces the size of the pupil. This is
the pupillary light reflex, which is an important test of brainstem function. Furthermore, the pupil will
dilate if a person sees an object of interest.
The pupil gets wider in the dark but narrower in light. When narrow, the diameter is 2 to 4
millimeters. In the dark it will be the same at first, but will approach the maximum distance for a wide
pupil 3 to 8 mm. In any human age group there is however considerable variation in maximal pupil
size. For example, at the peak age of 15, the dark-adapted pupil can vary from 4 mm to 9 mm with

different individuals. After 25 years of age the average pupil size decreases, though not at a steady
rate.[3][4] At this stage the pupils do not remain completely still, therefore may lead to oscillation, which
may intensify and become known as hippus. The constriction of the pupil and near vision are closely
tied. In bright light, the pupils constrict to prevent aberrations of light rays and thus attain their
expected acuity; in the dark this is not necessary, so it is chiefly concerned with admitting sufficient
light into the eye.[5]
A condition called bene dilitatism occurs when the optic nerves are partially damaged. This condition
is typified by chronically widened pupils due to the decreased ability of the optic nerves to respond to
light. In normal lighting, people afflicted with this condition normally have dilated pupils, and bright
lighting can cause pain. At the other end of the spectrum, people with this condition have trouble
seeing in darkness. It is necessary for these people to be especially careful when driving at night
due to their inability to see objects in their full perspective. This condition is not otherwise dangerous.

Effect of drugs[edit]
The sphincter muscle has a parasympathetic innervation, and the dilator has a sympathetic
innervation. In pupillary constriction induced by pilocarpine, not only is the sphincter nerve supply
activated but that of the dilator is inhibited. The reverse is true, so control of pupil size is controlled
by differences in contraction intensity of each muscle.
Certain drugs cause constriction of the pupils, such as alcohol and opioids.[citation needed] Other drugs,
such as atropine, LSD, MDMA,mescaline, psilocybin mushrooms, cocaine and amphetamines may
cause pupil dilation.[citation needed]
Another term for the constriction of the pupil is miosis. Substances that cause miosis are described
as miotic. Dilation of the pupil ismydriasis. Dilation can be caused by mydriatic substances such as
an eye drop solution containing tropicamide.

Other animals[edit]
The W-shaped pupil of the cuttlefish expanding when the lights are turned off

Not all animals have circular pupils. Some have slits or ovals which may be oriented vertically, as
in crocodiles, vipers, cats and foxes, or horizontally as in some rays, flying
frogs, mongooses andartiodactyls such as sheep, elk, red deer, reindeer and hippopotamus, as well
as the domestichorse. Goats, toads and octopus pupils tend to be horizontal and rectangular with
rounded corners. Some skates and rays have crescent shaped pupils, [6] gecko pupils range from
circular, to a slit, to a series of pinholes, and the cuttlefish pupil is a smoothly curving W shape.
There may be differences in pupil shape even between closely related animals. In felids, there are
differences between small- and large eyed species. The domestic cat (Felis sylvestris
domesticus) has vertical slit pupils, its large relative the Siberian tiger (Panthera tigris altaica) has

circular pupils and the Eurasian lynx (Lynx lynx) is intermediate between those of the domestic cat
and the Siberian tiger. A similar difference between small and large species may be present in
canines. The small European red fox(Vulpes vulpes) has vertical slit pupils whereas their large
relatives, the gray wolf (Canis lupus lupus) and domestic dogs (Canis lupus familiaris) have round
pupils.
One explanation for the evolution of slit pupils is that they can exclude light more effectively than a
circular pupil. This would explain why slit pupils tend to be found in the eyes of animals with
a crepuscular or nocturnal lifestyle that need to protect their eyes during daylight. Constriction of a
circular pupil (by a ring-shaped muscle) is less complete than closure of a slit pupil, which uses two
additional muscles that laterally compress the pupil. [7] For example, the cat's slit pupil can change the
light intensity on the retina 135-fold compared to 10-fold in humans.[8] However, this explanation does
not account for circular pupils that can be closed to a very small size (e.g., 0.5 mm in the tarsier) and
the rectangular pupils of many ungulates which do not close to a narrow slit in bright light. [9] An
alternative explanation is that a partially constricted circular pupil shades the peripheral zones of the
lens which would lead to poorly focused images at relevant wavelengths. The vertical slit pupil
allows for use of all wavelengths across the full diameter of the lens, even in bright light. [2] It has also
been suggested that in ambush predators such as some snakes, vertical slit pupils may aid in
camouflage, breaking up the circular outline of the eye. [10]
In a study of Australian snakes, pupil shapes correlated both with diel activity times and with foraging
behaviour. Most snake species with vertical pupils were nocturnal and also ambush foragers, and
most snakes with circular pupils were diurnal and active foragers. Overall, foraging behaviour
predicted pupil shape accurately in more cases than did diel time of activity, because many activeforaging snakes with circular pupils were not diurnal. It has been suggested that there may be a
similar link between foraging behaviour and pupil shape amongst the felidae and canidae discussed
above.[10]
Since any pupil provides better image quality for contours perpendicular to the pupils long axis,
[11]

vertical slits improve the horizontal resolution, which is important for stereopsis, and are therefore

common in predators. This advantage vanishes when there is little or no overlap between the two
eyes, as in prey animals. For those animals, vertical resolution is generally more important, as when
detecting an approaching predator, or gauging distance for a jump. [original research?]