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The nictitating membrane (from Latin '' nictare'', to blink) is a transparent or
translucent In the field of optics, transparency (also called pellucidity or diaphaneity) is the physical property of allowing light to pass through the material without appreciable light scattering by particles, scattering of light. On a macroscopic scale ...
third
eyelid An eyelid ( ) is a thin fold of skin that covers and protects an eye. The levator palpebrae superioris muscle retracts the eyelid, exposing the cornea to the outside, giving vision. This can be either voluntarily or involuntarily. "Palpebral ...
present in some animals that can be drawn across the
eye An eye is a sensory organ that allows an organism to perceive visual information. It detects light and converts it into electro-chemical impulses in neurons (neurones). It is part of an organism's visual system. In higher organisms, the ey ...
from the
medial Medial may refer to: Mathematics * Medial magma, a mathematical identity in algebra Geometry * Medial axis, in geometry the set of all points having more than one closest point on an object's boundary * Medial graph, another graph that repr ...
canthus The canthus (: canthi, palpebral commissures) is either corner of the eye where the upper and lower eyelids meet. More specifically, the inner and outer canthi are, respectively, the medial and lateral ends/angles of the palpebral fissure. The ...
to protect and moisten it while maintaining vision. Most Anura (tailless amphibians), some reptiles, birds, and
shark Sharks are a group of elasmobranch cartilaginous fish characterized by a ribless endoskeleton, dermal denticles, five to seven gill slits on each side, and pectoral fins that are not fused to the head. Modern sharks are classified within the ...
s, and some mammals (such as
cat The cat (''Felis catus''), also referred to as the domestic cat or house cat, is a small domesticated carnivorous mammal. It is the only domesticated species of the family Felidae. Advances in archaeology and genetics have shown that the ...
s,
beaver Beavers (genus ''Castor'') are large, semiaquatic rodents of the Northern Hemisphere. There are two existing species: the North American beaver (''Castor canadensis'') and the Eurasian beaver (''C. fiber''). Beavers are the second-large ...
s,
polar bear The polar bear (''Ursus maritimus'') is a large bear native to the Arctic and nearby areas. It is closely related to the brown bear, and the two species can Hybrid (biology), interbreed. The polar bear is the largest extant species of bear ...
s, seals and
aardvark Aardvarks ( ; ''Orycteropus afer'') are medium-sized, burrowing, nocturnal mammals native to Africa. Aardvarks are the only living species of the family Orycteropodidae and the order Tubulidentata. They have a long proboscis, similar to a pi ...
s) have full nictitating membranes; in many other
mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s, a small,
vestigial Vestigiality is the retention, during the process of evolution, of genetically determined structures or attributes that have lost some or all of the ancestral function in a given species. Assessment of the vestigiality must generally rely on co ...
portion of the nictitating membrane remains in the corner of the eye. It is often informally called a third eyelid or haw; the scientific terms for it are the ''plica semilunaris'', ''membrana nictitans'', or ''palpebra tertia''.


Description

The nictitating membrane is a transparent or
translucent In the field of optics, transparency (also called pellucidity or diaphaneity) is the physical property of allowing light to pass through the material without appreciable light scattering by particles, scattering of light. On a macroscopic scale ...
third
eyelid An eyelid ( ) is a thin fold of skin that covers and protects an eye. The levator palpebrae superioris muscle retracts the eyelid, exposing the cornea to the outside, giving vision. This can be either voluntarily or involuntarily. "Palpebral ...
present in some animals that can be drawn across the
eye An eye is a sensory organ that allows an organism to perceive visual information. It detects light and converts it into electro-chemical impulses in neurons (neurones). It is part of an organism's visual system. In higher organisms, the ey ...
for protection and to moisten it while maintaining vision. The term comes from the Latin word '' nictare'', meaning "to blink". It is often called a ''third eyelid'' or ''haw'', and may be referred to in scientific terminology as the ''plica semilunaris'', ''membrana nictitans'', or ''palpebra tertia''. Unlike the upper and lower
eyelid An eyelid ( ) is a thin fold of skin that covers and protects an eye. The levator palpebrae superioris muscle retracts the eyelid, exposing the cornea to the outside, giving vision. This can be either voluntarily or involuntarily. "Palpebral ...
s, the nictitating membrane moves horizontally across the eyeball. In many species, any stimulus to the eyeball (such as a puff of air) will result in reflex nictitating membrane response. This reflex is widely used as the basis for experiments on
classical conditioning Classical conditioning (also respondent conditioning and Pavlovian conditioning) is a behavioral procedure in which a biologically potent Stimulus (physiology), stimulus (e.g. food, a puff of air on the eye, a potential rival) is paired with a n ...
in rabbits.


Distribution

Fully developed nictitating membranes are found in
fish A fish (: fish or fishes) is an aquatic animal, aquatic, Anamniotes, anamniotic, gill-bearing vertebrate animal with swimming fish fin, fins and craniate, a hard skull, but lacking limb (anatomy), limbs with digit (anatomy), digits. Fish can ...
,
amphibian Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...
s,
reptile Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...
s,
bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
s and mammals, but are rare in
primate Primates is an order (biology), order of mammals, which is further divided into the Strepsirrhini, strepsirrhines, which include lemurs, galagos, and Lorisidae, lorisids; and the Haplorhini, haplorhines, which include Tarsiiformes, tarsiers a ...
s. In humans, the plica semilunaris (also known as the semilunar fold) and its associated muscles are homologous to the nictitating membranes seen in some other mammals and other vertebrates. In most primate species, a plica semilunaris is generally not present, although fully developed nictitating membranes can be found in
lemur Lemurs ( ; from Latin ) are Strepsirrhini, wet-nosed primates of the Superfamily (biology), superfamily Lemuroidea ( ), divided into 8 Family (biology), families and consisting of 15 genera and around 100 existing species. They are Endemism, ...
s and lorisoid primates. Some mammals, such as
camel A camel (from and () from Ancient Semitic: ''gāmāl'') is an even-toed ungulate in the genus ''Camelus'' that bears distinctive fatty deposits known as "humps" on its back. Camels have long been domesticated and, as livestock, they provid ...
s,
polar bear The polar bear (''Ursus maritimus'') is a large bear native to the Arctic and nearby areas. It is closely related to the brown bear, and the two species can Hybrid (biology), interbreed. The polar bear is the largest extant species of bear ...
s, seals and
aardvark Aardvarks ( ; ''Orycteropus afer'') are medium-sized, burrowing, nocturnal mammals native to Africa. Aardvarks are the only living species of the family Orycteropodidae and the order Tubulidentata. They have a long proboscis, similar to a pi ...
s, have full nictitating membranes, and many mammals retain a small,
vestigial Vestigiality is the retention, during the process of evolution, of genetically determined structures or attributes that have lost some or all of the ancestral function in a given species. Assessment of the vestigiality must generally rely on co ...
portion of the membrane in the corner of the eye. A gland of the third eyelid (nictitans gland) or Harder's gland is attached to the nictating membranes of some animals and may produce up to 50% of the
tear film Tear, tears or tearing may refer to: * Tearing, the act of breaking apart a material by force * Tears, a clear liquid secreted by the tear gland in the eyes of land mammals Arts and entertainment Literature * "Tears", a poem by Edward Thomas ...
.


Functions

The nictitating membrane is normally translucent. In some diving animals, including
sea lion Sea lions are pinnipeds characterized by external ear flaps, long foreflippers, the ability to walk on all fours, short and thick hair, and a big chest and belly. Together with the fur seals, they make up the family Otariidae, eared seals. ...
s, it is activated on land, to remove sand and other debris—its function in most animals. In
crocodiles Crocodiles (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 more loosely to include all extant member ...
, it protects their eyes from water but also hinders their focus under water. In some diving animals, for example
beaver Beavers (genus ''Castor'') are large, semiaquatic rodents of the Northern Hemisphere. There are two existing species: the North American beaver (''Castor canadensis'') and the Eurasian beaver (''C. fiber''). Beavers are the second-large ...
s and
manatee Manatees (, family (biology), family Trichechidae, genus ''Trichechus'') are large, fully aquatic, mostly herbivory, herbivorous marine mammals sometimes known as sea cows. There are three accepted living species of Trichechidae, representing t ...
s, it is transparent and moves across the eye to protect it while under water. Birds can actively control their nictitating membrane. In birds of prey, the membrane also serves to protect the parents' eyes from their chicks while they are feeding them, and when
peregrine falcon The peregrine falcon (''Falco peregrinus''), also known simply as the peregrine, is a Cosmopolitan distribution, cosmopolitan bird of prey (raptor) in the family (biology), family Falconidae renowned for its speed. A large, Corvus (genus), cro ...
s go into their dives, they will blink repeatedly with their nictitating membranes to clear debris and spread moisture across the eyes.
Woodpecker Woodpeckers are part of the bird family (biology), family Picidae, which also includes the piculets, wrynecks and sapsuckers. Members of this family are found worldwide, except for Australia, New Guinea, New Zealand, Madagascar and the extreme ...
s tighten their nictitating membrane a millisecond prior to their beak impacting the trunk of a tree to prevent shaking-induced retinal injury.Wygnanski-Jaffe T, Murphy CJ, Smith C, Kubai M, Christopherson P, Ethier CR, Levin AV. (2007) Protective ocular mechanisms in woodpeckers ''Eye'' 21, 83–89. The nictitating membrane can be used to protect the eye while attacking prey, as in
shark Sharks are a group of elasmobranch cartilaginous fish characterized by a ribless endoskeleton, dermal denticles, five to seven gill slits on each side, and pectoral fins that are not fused to the head. Modern sharks are classified within the ...
s. It can also protect the eye from
ultraviolet radiation Ultraviolet radiation, also known as simply UV, is electromagnetic radiation of wavelengths of 10–400 nanometers, shorter than that of visible light, but longer than X-rays. UV radiation is present in sunlight and constitutes about 10% of t ...
, similar to its role in polar bears to prevent
snow blindness Photokeratitis or ultraviolet keratitis is a painful eye condition caused by exposure of insufficiently protected Human eye, eyes to the ultraviolet (UV) rays from either natural (e.g. intense direct or reflected sunlight) or artificial (e.g. t ...
.


Vestigiality

Nictitating membranes in cats and dogs do not have many muscle fibers, so they are not usually visible; chronic visibility should be taken as a sign of poor condition or ill health. The membrane can, however, be seen clearly by gently opening the eye of the healthy animal when it is asleep, or by pushing down/applying pressure on the eyeball, which will cause it to appear. In some breeds of dogs, the nictitating membrane can be prone to
prolapse In medicine, prolapse is a condition in which organ (anatomy), organs fall down or slip out of place. It is used for organs protruding through the vagina, rectum, or for the misalignment of the valves of the heart. A spinal disc herniation is al ...
of the gland of the third eyelid, resulting in a condition called cherry eye.Artem Cheprasov
Why do cats have an extra eyelid?
Guru Magazine. 7 Feb 2013. (Accessed 26 Mar 2013)


See also

* Accessory abducens nucleus *
Human vestigiality In the context of human evolution, vestigiality involves those phenotypic trait, traits occurring in humans that have lost all or most of their original function (biology), function through evolution. Although structures called vestigial often a ...


Notes


References


External links

* {{youTube, dgh6ABfdlAU, Hawk, transparent eyelids (nicitating membrane slow motion video) Eye Vertebrate anatomy fi:Vilkkuluomi