Nictitating membrane

Last updated
The nictitating membrane of a masked lapwing as it closes over the left eye, originating from the medial canthus Bird blink-edit.jpg
The nictitating membrane of a masked lapwing as it closes over the left eye, originating from the medial canthus

The nictitating membrane (from Latin nictare , to blink) is a transparent or translucent third eyelid present in some animals that can be drawn across the eye from the medial canthus to protect and moisten it while maintaining vision. Most Anura [1] [a] (tailless amphibians), some reptiles, birds, and sharks, and some mammals (such as cats, beavers, polar bears, seals and aardvarks) have full nictitating membranes; in many other mammals, a small, vestigial 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.

Contents

Description

Nictating membrane of a blue shark.tiff
Haliaeetus leucocephalus LC0198.jpg
Third eyelid on a domestic dog.jpg
Common Goldeneye showing nictitating membrane - crop.JPG
Nictitating membranes of, top to bottom and left to right, a blue shark, a bald eagle (mid-blink), a domestic dog, a common goldeneye, a cat, and a common buzzard (deployed asynchronously on left and right eye).

The nictitating membrane is a transparent or translucent third eyelid present in some animals that can be drawn across the eye 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 eyelids, 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 in rabbits. [3]

Distribution

Fully developed nictitating membranes are found in fish, amphibians, reptiles, birds and mammals, but are rare in primates. [4] [5] 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. [6] In most primate species, a plica semilunaris is generally not present, although fully developed nictitating membranes can be found in lemurs and lorisoid primates. [7] [8] Some mammals, such as camels, polar bears, seals and aardvarks, have full nictitating membranes, and many mammals retain a small, vestigial 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. [9]

Functions

The nictitating membrane is normally translucent. In some diving animals, including sea lions, it is activated on land, to remove sand and other debris—its function in most animals. In crocodiles, it protects their eyes from water but also hinders their focus under water. In some diving animals, for example beavers and manatees, it is transparent and moves across the eye to protect it while under water.

Birds can actively control their nictitating membrane. [10] In birds of prey, the membrane also serves to protect the parents' eyes from their chicks while they are feeding them, and when peregrine falcons go into their 200-mile-per-hour (320 km/h) dives, they will blink repeatedly with their nictitating membranes to clear debris and spread moisture across the eyes. Woodpeckers tighten their nictitating membrane a millisecond prior to their beak impacting the trunk of a tree to prevent shaking-induced retinal injury. [11]

The nictitating membrane can be used to protect the eye while attacking prey, as in sharks.

It can also protect the eye from ultraviolet radiation, similar to its role in polar bears to prevent snow blindness.

Vestigiality

The plica semilunaris of conjunctiva is a vestigial remnant of a nictitating membrane in humans. Gray892.png
The plica semilunaris of conjunctiva is a vestigial remnant of a nictitating membrane in humans.

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 of the gland of the third eyelid, resulting in a condition called cherry eye. [9]

See also

Notes

  1. Except Hymenochirus , Pseudhymenochirus and Pipa species [2] :439

Related Research Articles

<span class="mw-page-title-main">Tears</span> Clear liquid secreted from glands in eyes of mammals

Tears are a clear liquid secreted by the lacrimal glands found in the eyes of all land mammals. Tears are made up of water, electrolytes, proteins, lipids, and mucins that form layers on the surface of eyes. The different types of tears—basal, reflex, and emotional—vary significantly in composition.

<span class="mw-page-title-main">Vestigiality</span> Evolutionary retention of no longer needed structures in living organisms

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 comparison with homologous features in related species. The emergence of vestigiality occurs by normal evolutionary processes, typically by loss of function of a feature that is no longer subject to positive selection pressures when it loses its value in a changing environment. The feature may be selected against more urgently when its function becomes definitively harmful, but if the lack of the feature provides no advantage, and its presence provides no disadvantage, the feature may not be phased out by natural selection and persist across species.

Haw or HAW may refer to:

Blinking is a bodily function; it is a semi-autonomic rapid closing of the eyelid. A single blink is determined by the forceful closing of the eyelid or inactivation of the levator palpebrae superioris and the activation of the palpebral portion of the orbicularis oculi, not the full open and close. It is an essential function of the eye that helps spread tears across and remove irritants from the surface of the cornea and conjunctiva.

<span class="mw-page-title-main">Eyelid</span> Thin fold of skin that covers and protects the eye

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" means relating to the eyelids. Its key function is to regularly spread the tears and other secretions on the eye surface to keep it moist, since the cornea must be continuously moist. They keep the eyes from drying out when asleep. Moreover, the blink reflex protects the eye from foreign bodies. A set of specialized hairs known as lashes grow from the upper and lower eyelid margins to further protect the eye from dust and debris.

<span class="mw-page-title-main">Cherry eye</span> Prolapse of the tear gland of the third eyelid in animals

Cherry eye is a disorder of the nictitating membrane (NM), also called the third eyelid, present in the eyes of dogs and cats. Cherry eye is most often seen in young dogs under the age of two. Common misnomers include adenitis, hyperplasia, adenoma of the gland of the third eyelid; however, cherry eye is not caused by hyperplasia, neoplasia, or primary inflammation. In many species, the third eyelid plays an essential role in vision by supplying oxygen and nutrients to the eye via tear production. Normally, the gland can turn inside-out without detachment. Cherry eye results from a defect in the retinaculum which is responsible for anchoring the gland to the periorbita. This defect causes the gland to prolapse and protrude from the eye as a red fleshy mass. Problems arise as sensitive tissue dries out and is subjected to external trauma Exposure of the tissue often results in secondary inflammation, swelling, or infection. If left untreated, this condition can lead to dry eye syndrome and other complications.

<span class="mw-page-title-main">Eyelash</span> Facial feature

An eyelash is one of the hairs that grows at the edges of the top and bottom eyelids, spanning outwards and away from the eyes. The lashes grow in up to six layers on each of the upper and lower eyelids. Eyelashes serve to protect the eye from debris, dust, and small particles, and are highly sensitive to touch, thus providing a warning that an object is near the eye, which then reflexively closes or flutters to rid the area of the object. The eyelid margin from which lashes grow is among the most sensitive parts of the human body, with many nerve endings enveloping the roots of the lashes, giving it sensitivity to very light tactile input even at the tips of the lashes, enabling it to trigger the blink reflex when touched. Eyelashes are also an important component of physical attractiveness, with long prominent lashes giving the illusion of large, gazing eyes, and drawing attention to the eyes.

<span class="mw-page-title-main">Plica semilunaris of conjunctiva</span> Small fold of mucous membrane in the eye

The plica semilunaris is a small fold of bulbar conjunctiva on the medial canthus of the eye. It functions during movement of the eye, to help maintain tear drainage via the lacrimal lake, and to permit greater rotation of the globe, for without the plica the conjunctiva would attach directly to the eyeball, restricting movement. It is the vestigial remnant of the nictitating membrane which is drawn across the eye for protection, and is present in other animals such as birds, reptiles, and fish, but is rare in mammals, mainly found in monotremes and marsupials. Its associated muscles are also vestigial. It is loose, thus eye movements are not restricted by it. Only one species of primate, the Calabar angwantibo, is known to have a functioning nictitating membrane.

<span class="mw-page-title-main">Brille</span> Part of the anatomy of the eye in some animals

The brille is the layer of transparent, immovable disc-shaped skin or scale covering the eyes of some animals for protection, especially in animals without eyelids. In squamate reptiles both the nictitating membrane and the eyelids have been suggested as the evolutionary origin of the brille, but embryonic studies supports the fusion of the dorsal and ventral eyelids. Brille means "spectacles" or "glasses" in German, Norwegian, and Danish.

<span class="mw-page-title-main">Cat senses</span> Senses of Felis catus

Cat senses are adaptations that allow cats to be highly efficient predators. Cats are good at detecting movement in low light, have an acute sense of hearing and smell, and their sense of touch is enhanced by long whiskers that protrude from their heads and bodies. These senses evolved to allow cats to hunt effectively at dawn and dusk.

<span class="mw-page-title-main">Human vestigiality</span> Human traits which lost their original function through evolution

In the context of human evolution, vestigiality involves those traits occurring in humans that have lost all or most of their original function through evolution. Although structures called vestigial often appear functionless, a vestigial structure may retain lesser functions or develop minor new ones. In some cases, structures once identified as vestigial simply had an unrecognized function. Vestigial organs are sometimes called rudimentary organs. Many human characteristics are also vestigial in other primates and related animals.

<span class="mw-page-title-main">Equine vision</span> Eyesight capabilities of horses

The equine eye is one of the largest of any land mammal. Its visual abilities are directly related to the animal's behavior; for example, it is active during both day and night, and it is a prey animal. Both the strengths and weaknesses of the horse's visual abilities should be taken into consideration when training the animal, as an understanding of the horse's eye can help to discover why the animal behaves the way it does in various situations.

<span class="mw-page-title-main">Xerocole</span> Any animal adapted to live in the desert

A xerocole, is a general term referring to any animal that is adapted to live in a desert. The main challenges xerocoles must overcome are lack of water and excessive heat. To conserve water they avoid evaporation and concentrate excretions. Some are so adept at conserving water or obtaining it from food that they do not need to drink at all. To escape the desert heat, xerocoles tend to be either nocturnal or crepuscular.

<span class="mw-page-title-main">Mammalian eye</span>

Mammals normally have a pair of eyes. Although mammalian vision is not so excellent as bird vision, it is at least dichromatic for most of mammalian species, with certain families possessing a trichromatic color perception.

<span class="mw-page-title-main">Blindness in animals</span> Animals with limited visual perception

Visual perception in animals plays an important role in the animal kingdom, most importantly for the identification of food sources and avoidance of predators. For this reason, blindness in animals is a unique topic of study.

<i>Thelazia</i> Genus of roundworms

Thelazia is a genus of nematode worms which parasitize the eyes and associated tissues of various bird and mammal hosts, including humans. They are often called "eyeworms", and infestation with Thelazia species is referred to as "thelaziasis". Adults are usually found in the eyelids, tear glands, tear ducts, or the so-called "third eyelid". Occasionally, they are found in the eyeball itself, either under the conjunctiva or in the vitreous cavity of the eyeball. All species of Thelazia for which the life cycle has been studied are transmitted by species of Diptera (flies) which do not bite, but which feed on tears.

<span class="mw-page-title-main">Thelaziasis</span> Medical condition

Thelaziasis is the term for infestation with parasitic nematodes of the genus Thelazia. The adults of all Thelazia species discovered so far inhabit the eyes and associated tissues of various mammal and bird hosts, including humans. Thelazia nematodes are often referred to as "eyeworms".

<span class="mw-page-title-main">Harderian gland</span> Gland in the eye socket of tetrapods

The Harderian gland is a gland found within the eye's orbit that occurs in tetrapods that possess a nictitating membrane.

<span class="mw-page-title-main">Sublingua</span> Secondary tongue found in some primates

The sublingua ("under-tongue") is a muscular secondary tongue found below the primary tongue in tarsiers and living strepsirrhine primates, which includes lemurs and lorisoids. Although it is most fully developed in these primates, similar structures can be found in some other mammals, such as marsupials, treeshrews, and colugos. This "second tongue" lacks taste buds, and in lemuriforms, it is thought to be used to remove hair and other debris from the toothcomb, a specialized dental structure used to comb the fur during oral grooming.

<span class="mw-page-title-main">Eagle eye</span>

The eagle eye is among the sharpest in the animal kingdom, with an eyesight estimated at 4 to 8 times stronger than that of the average human. Although an eagle may only weigh 4.5 kilograms (10 lb), its eyes are roughly the same size as those of a human. Eagle weight varies: a small eagle could weigh 700 grams (1.5 lb), while a larger one could weigh 6.5 kilograms (14 lb); an eagle of about 4.5 kilograms (9.9 lb) weight could have eyes as big as that of a human who weighs 91 kilograms (200 lb). Although the size of the eagle eye is about the same as that of a human being, the back side shape of the eagle eye is flatter. Their eyes are stated to be larger than their brain, by weight. Color vision with resolution and clarity are the most prominent features of eagles' eyes, hence sharp-sighted people are sometimes referred to as "eagle-eyed". Eagles can identify five distinctly colored squirrels and locate their prey even if hidden.

References

  1. Rolleston, George; Jackson, William Hatchett (1898). Forms of Animal Life : A Manual of Comparative Anatomy, with Descriptions of Selected Types. Clarendon Press. p. 401.
  2. Vitt, Laurie J.; Caldwell, Janalee P. (2009). "Chapter 17 – Frogs". Herpetology: An Introductory Biology of Amphibians and Reptiles (Third ed.). Academic Press. pp. 435–482. doi:10.1016/B978-0-12-374346-6.00017-1. ISBN   978-0-12-374346-6.
  3. Gormezano, I. N. Schneiderman, E. Deaux, and I. Fuentes (1962) Nictitating Membrane: Classical Conditioning and Extinction in the Albino Rabbit Science138:33–34.
  4. Butler, Ann B.; Hodos, William (2 September 2005). Comparative Vertebrate Neuroanatomy: Evolution and Adaptation . John Wiley & Sons. p.  215. ISBN   978-0-471-73383-6.
  5. Paul Miller, Why do cats have an inner eyelid as well as outer ones? Scientific American. 20 Nov 2006. (Accessed 2 Nov 2011)
  6. The Eye: Basic Sciences in Practice by John V. Forrester, p. 82
  7. Osman Hill, W. C. (1953). Primates Comparative Anatomy and Taxonomy I—Strepsirhini. Edinburgh Univ Pubs Science & Maths, No 3. Edinburgh University Press. p. 13. OCLC   500576914.
  8. Ankel-Simons, F. (2007). Primate Anatomy (3rd ed.). Academic Press. p. 471. ISBN   978-0-12-372576-9.
  9. 1 2 Artem Cheprasov, Why do cats have an extra eyelid? Archived 2016-01-01 at the Wayback Machine Guru Magazine. 7 Feb 2013. (Accessed 26 Mar 2013)
  10. Frans C. Stades, Milton Wyman, Michael H. Boevé, Willy Neumann, Bernhard Spiess. Ophthalmology for the Veterinary Practitioner. 105–106
  11. 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.
Listen to this article (4 minutes)
Sound-icon.svg
This audio file was created from a revision of this article dated 25 September 2019 (2019-09-25), and does not reflect subsequent edits.