The pupil is a black hole located in the center of the iris of the

eye Eyes are organs of the visual system. They provide living organisms with vision, the ability to receive and process visual detail, as well as enabling several photo response functions that are independent of vision. Eyes detect light and conv ...

that allows light to strike the retina.Cassin, B. and Solomon, S. (1990) ''Dictionary of Eye Terminology''. Gainesville, Florida: Triad Publishing Company. It appears black because light rays entering the pupil are either absorbed by the tissues inside the eye directly, or absorbed after diffuse reflections within the eye that mostly miss exiting the narrow pupil. The term "pupil" was coined by Gerard of Cremona. In humans, the pupil is round, but its shape varies between species; some

cats The cat (''Felis catus'') is a domestic species of small carnivorous mammal. It is the only domesticated species in the family Felidae and is commonly referred to as the domestic cat or house cat to distinguish it from the wild members of t ...

reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...

s, and

fox Foxes are small to medium-sized, omnivorous mammals belonging to several genera of the family Canidae. They have a flattened skull, upright, triangular ears, a pointed, slightly upturned snout, and a long bushy tail (or ''brush''). Twelv ...

es have vertical slit pupils, goats have horizontally oriented pupils, and some catfish have annular types. 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 In an optical system, the entrance pupil is the optical image of the physical aperture stop, as 'seen' through the front (the object side) of the lens system. The corresponding image of the aperture as seen through the back of the lens system is ...

, which does not exactly correspond to the location and size of the physical pupil because it is magnified by the cornea. On the inner edge lies a prominent structure, the collarette, marking the junction of the embryonic pupillary membrane covering the embryonic pupil.

Structure

The pupil is a hole located in the center of the iris of the

that allows light to strike the retina. It appears black because light rays entering the pupil are either absorbed by the tissues inside the eye directly, or absorbed after diffuse reflections within the eye that mostly miss exiting the narrow pupil.

Function

The iris is a contractile structure, consisting mainly of

smooth muscle Smooth muscle is an involuntary non-striated muscle, so-called because it has no sarcomeres and therefore no striations (''bands'' or ''stripes''). It is divided into two subgroups, single-unit and multiunit smooth muscle. Within single-unit mus ...

, 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. This is known as the pupillary light reflex. The iris contains two groups of smooth muscles; a circular group called the sphincter pupillae, and a radial group called the

dilator pupillae The iris dilator muscle (pupil dilator muscle, pupillary dilator, radial muscle of iris, radiating fibers), is a smooth muscle of the eye, running radially in the iris and therefore fit as a dilator. The pupillary dilator consists of a spokelike ...

. 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.

Effect of light

The pupil gets wider in the dark and 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. However, in any human age group there is 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. At this stage the pupils do not remain completely still, therefore may lead to oscillation, which may intensify and become known as

hippus Pupillary hippus, also known as pupillary athetosis, is spasmodic, rhythmic, but regular dilating and contracting pupillary movements between the sphincter and dilator muscles.Cassin, B. and Solomon, S. ''Dictionary of Eye Terminology''. Gainesvill ...

. 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. When bright light is shone on the eye, light-sensitive cells in the retina, including rod and cone photoreceptors and

melanopsin Melanopsin is a type of photopigment belonging to a larger family of light-sensitive retinal proteins called opsins and encoded by the gene ''Opn4''. In the mammalian retina, there are two additional categories of opsins, both involved in the for ...

ganglion cells, will send signals to the

oculomotor nerve The oculomotor nerve, also known as the third cranial nerve, cranial nerve III, or simply CN III, is a cranial nerve that enters the orbit through the superior orbital fissure and innervates extraocular muscles that enable most movements of ...

, specifically the

parasympathetic The parasympathetic nervous system (PSNS) is one of the three divisions of the autonomic nervous system, the others being the sympathetic nervous system and the enteric nervous system. The enteric nervous system is sometimes considered part of t ...

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 The brainstem (or brain stem) is the posterior stalk-like part of the brain that connects the cerebrum with the spinal cord. In the human brain the brainstem is composed of the midbrain, the pons, and the medulla oblongata. The midbrain is cont ...

function. Furthermore, the pupil will dilate if a person sees an object of interest.

Clinical significance

Effect of drugs

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. Certain drugs cause constriction of the pupils, such as

opioids Opioids are substances that act on opioid receptors to produce morphine-like effects. Medically they are primarily used for pain relief, including anesthesia. Other medical uses include suppression of diarrhea, replacement therapy for opioid use ...

. Other drugs, such as

atropine Atropine is a tropane alkaloid and anticholinergic medication used to treat certain types of nerve agent and pesticide poisonings as well as some types of slow heart rate, and to decrease saliva production during surgery. It is typically given i ...

, LSD, MDMA,

mescaline Mescaline or mescalin (3,4,5-trimethoxyphenethylamine) is a naturally occurring psychedelic protoalkaloid of the substituted phenethylamine class, known for its hallucinogenic effects comparable to those of LSD and psilocybin. Biological sou ...

, psilocybin mushrooms, cocaine and amphetamines may cause pupil dilation. 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. Another term for the constriction of the pupil is miosis. Substances that cause miosis are described as miotic. Dilation of the pupil is

mydriasis Mydriasis is the dilation of the pupil, usually having a non-physiological cause, or sometimes a physiological pupillary response. Non-physiological causes of mydriasis include disease, trauma, or the use of certain types of drugs. Normally, as ...

. Dilation can be caused by mydriatic substances such as an eye drop solution containing tropicamide.

Diseases

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 affected by 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.

Size

The size of the pupil (often measured as diameter) can be a

symptom Signs and symptoms are the observed or detectable signs, and experienced symptoms of an illness, injury, or condition. A sign for example may be a higher or lower temperature than normal, raised or lowered blood pressure or an abnormality showin ...

of an underlying disease. Dilation of the pupil is known as

and contraction as miosis. Not all variations in size are indicative of disease however. In addition to dilation and contraction caused by light and darkness, it has been shown that solving simple multiplication problems affects the size of the pupil. The simple act of recollection can dilate the size of the pupil, however when the brain is required to process at a rate above its maximum capacity, the pupils contract. There is also evidence that pupil size is related to the extent of positive or negative emotional arousal experienced by a person.

Animals

Not all animals have circular pupils. Some have slits or ovals which may be oriented vertically, as in

crocodile Crocodiles (family (biology), family Crocodylidae) or true crocodiles are large semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term crocodile is sometimes used even more loosely to inclu ...

s, vipers,

and

es, or horizontally as in some rays,

flying frog A flying frog (also called a gliding frog) is a frog that has the ability to achieve gliding flight. This means it can descend at an angle less than 45° relative to the horizontal. Other nonflying arboreal frogs can also descend, but only at angle ...

s, mongooses and artiodactyls such as sheep,

elk The elk (''Cervus canadensis''), also known as the wapiti, is one of the largest species within the deer family, Cervidae, and one of the largest terrestrial mammals in its native range of North America and Central and East Asia. The common ...

, red deer, reindeer and hippopotamus, as well as the domestic horse. Goats, toads and

octopus An octopus ( : octopuses or octopodes, see below for variants) is a soft-bodied, eight- limbed mollusc of the order Octopoda (, ). The order consists of some 300 species and is grouped within the class Cephalopoda with squids, cuttle ...

pupils tend to be horizontal and rectangular with rounded corners. Some skates and rays have crescent shaped pupils,

gecko Geckos are small, mostly carnivorous lizards that have a wide distribution, found on every continent except Antarctica. Belonging to the infraorder Gekkota, geckos are found in warm climates throughout the world. They range from . Geckos ar ...

pupils range from circular, to a slit, to a series of pinholes, and the

cuttlefish Cuttlefish or cuttles are marine molluscs of the order Sepiida. They belong to the class Cephalopoda which also includes squid, octopuses, and nautiluses. Cuttlefish have a unique internal shell, the cuttlebone, which is used for control of ...

pupil is a smoothly curving W shape. Although human pupils are normally circular, abnormalities like colobomas can result in unusual pupil shapes, such as teardrop, keyhole or oval pupil shapes. 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

red fox The red fox (''Vulpes vulpes'') is the largest of the true foxes and one of the most widely distributed members of the Order (biology), order Carnivora, being present across the entire Northern Hemisphere including most of North America, Europe ...

''(Vulpes vulpes)'' has vertical slit pupils whereas their large relatives, the

gray wolf The wolf (''Canis lupus''; : wolves), also known as the gray wolf or grey wolf, is a large canine native to Eurasia and North America. More than thirty subspecies of ''Canis lupus'' have been recognized, and gray wolves, as popularly ...

''(Canis lupus lupus)'' and domestic dogs ''(Canis lupus familiaris)'' have round pupils.

Evolution and adaptation

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 In zoology, a crepuscular animal is one that is active primarily during the twilight period, being matutinal, vespertine, or both. This is distinguished from diurnal and nocturnal behavior, where an animal is active during the hours of daylig ...

nocturnal Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sens ...

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. For example, the cat's slit pupil can change the light intensity on the retina 135-fold compared to 10-fold in humans. 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 Tarsiers ( ) are haplorhine primates of the family Tarsiidae, which is itself the lone extant family within the infraorder Tarsiiformes. Although the group was once more widespread, all of its species living today are found in Maritime Southeast ...

) and the rectangular pupils of many ungulates which do not close to a narrow slit in bright light. 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. 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.

Activity pattern and behavior

In a study of

Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...

n snakes, pupil shapes correlated both with diel activity times and with foraging behavior. Most snake species with vertical pupils were

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 active-foraging 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. A 2015 study confirmed the hypothesis that elongated pupils have increased dynamic range, and furthered the correlations with diel activity. However it noted that other hypotheses could not explain the orientation of the pupils. They showed that vertical pupils enable ambush predators to optimise their depth perception, and horizontal pupils to optimise the field of view and image quality of horizontal contours. They further explained why elongated pupils are correlated with the animal's height. File:Closeup of goat eye.jpg, A goat with horizontal rectangular pupils File:Taeniura_grabata_eye.jpg, A

stingray Stingrays are a group of sea rays, which are cartilaginous fish related to sharks. They are classified in the suborder Myliobatoidei of the order Myliobatiformes and consist of eight families: Hexatrygonidae (sixgill stingray), Plesiobatidae ( ...

with crescent pupils File:Crocodylus siamensis closeup.jpg, A

with thin vertical slit pupils File:Cuttlefish_eye.jpg, A

with W-shaped pupils File:Gecko-oeil.jpg, A

with 'thin string of pearls' pupils File:Catpupil03042006.jpg, A cat with thick vertical slit pupils

Society and culture

In a surprising number of unrelated languages, the etymological meaning of the term for pupil is "little person". This is true, for example, of the word ''pupil'' itself: this comes into English from Latin ''pūpilla'', which means "doll, girl", and is a diminutive form of ''pupa'', "girl". (The double meaning in Latin is preserved in English, where ''pupil'' means both "schoolchild" and "dark central portion of the eye within the iris".) This may be because the reflection of one's image in the pupil is a minuscule version of one's self.Human Universals and Human Culture, p4
In the Old Babylonian period (c. 1800-1600 BC) in ancient Mesopotamia, the expression "protective spirit of the eye" is attested, perhaps arising from the same phenomenon. The English phrase ''

apple of my eye The phrase "apple of my eye" refers in English to something or someone that one cherishes above all others. Originally, the phrase was simply an idiom referring to the pupil of the eye.

'' arises from an

Old English Old English (, ), or Anglo-Saxon, is the earliest recorded form of the English language, spoken in England and southern and eastern Scotland in the early Middle Ages. It was brought to Great Britain by Anglo-Saxon settlement of Britain, Anglo ...

usage, in which the word ''apple'' meant not only the fruit but also the pupil or eyeball.apple, n.
, '' Oxford English Dictionary Online'', 3rd ed. (Oxford University Press, 2008), § 6 B.

References

External links

* — "Sagittal Section Through the Eyeball" * — "Sagittal Section Through the Eyeball"
A pupil examination simulator
demonstrating the changes in pupil reactions for various nerve lesions. {{Authority control Ethology Articles containing video clips