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The sclera, also known as the white of the eye or, in older literature, as the tunica albuginea oculi, is the opaque, fibrous, protective, outer layer of the
human eye The human eye is a sensory organ, part of the sensory nervous system, that reacts to visible light and allows humans to use visual information for various purposes including seeing things, keeping balance, and maintaining circadian rhythm. ...
containing mainly
collagen Collagen () is the main structural protein in the extracellular matrix found in the body's various connective tissues. As the main component of connective tissue, it is the most abundant protein in mammals, making up from 25% to 35% of the whole ...
and some crucial
elastic fiber Elastic fibers (or yellow fibers) are an essential component of the extracellular matrix composed of bundles of proteins (elastin) which are produced by a number of different cell types including fibroblasts, endothelial, smooth muscle, and air ...
. In humans, and some other vertebrates, the whole sclera is white, contrasting with the coloured iris, but in most mammals, the visible part of the sclera matches the colour of the iris, so the white part does not normally show while other vertebrates have distinct colors for both of them. In the development of the
embryo An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male spe ...
, the sclera is derived from the neural crest. In children, it is thinner and shows some of the underlying pigment, appearing slightly blue. In the elderly, fatty deposits on the sclera can make it appear slightly yellow. People with dark skin can have naturally darkened sclerae, the result of melanin pigmentation. The human eye is relatively rare for having a pale sclera (relative to the iris). This makes it easier for one individual to identify where another individual is looking, and the
cooperative eye hypothesis The cooperative eye hypothesis is a proposed explanation for the appearance of the human eye. It suggests that the eye's distinctive visible characteristics evolved to make it easier for humans to follow another's gaze while communicating or while ...
suggests this has evolved as a method of nonverbal communication.


Structure

The sclera forms the posterior five-sixths of the
connective tissue Connective tissue is one of the four primary types of animal tissue, along with epithelial tissue, muscle tissue, and nervous tissue. It develops from the mesenchyme derived from the mesoderm the middle embryonic germ layer. Connective tiss ...
coat of the globe. It is continuous with the dura mater and the cornea, and maintains the shape of the globe, offering resistance to internal and external forces, and provides an attachment for the extraocular muscle insertions. The sclera is perforated by many nerves and vessels passing through the posterior scleral foramen, the hole that is formed by the optic nerve. At the
optic disc The optic disc or optic nerve head is the point of exit for ganglion cell axons leaving the eye. Because there are no rods or cones overlying the optic disc, it corresponds to a small blind spot in each eye. The ganglion cell axons form the ...
the outer two-thirds of the sclera continues with the dura mater (outer coat of the brain) via the dural sheath of the optic nerve. The inner third joins with some
choroid The choroid, also known as the choroidea or choroid coat, is a part of the uvea, the vascular layer of the eye, and contains connective tissues, and lies between the retina and the sclera. The human choroid is thickest at the far extreme rear ...
al tissue to form a plate ( lamina cribrosa) across the optic nerve with perforations through which the optic fibers (
fasciculi Fascicle or ''fasciculus'' may refer to: Anatomy and histology * Muscle fascicle, a bundle of skeletal muscle fibers * Nerve fascicle, a bundle of axons (nerve fibers) ** Superior longitudinal fasciculus *** Arcuate fasciculus ** Gracile fasci ...
) pass. The thickness of the sclera varies from 1 mm at the posterior pole to 0.3 mm just behind the
rectus muscle Rectus muscle (Latin: , "straight muscle") may refer to: In the trunk: * Rectus abdominis muscle In the eye: * Inferior rectus muscle * Lateral rectus muscle * Medial rectus muscle * Superior rectus muscle In the leg: * Rectus femoris muscle In ...
insertions. The sclera's blood vessels are mainly on the surface. Along with the vessels of the conjunctiva (which is a thin layer covering the sclera), those in the episclera render the inflamed eye bright red."eye, human."Encyclopædia Britannica from Encyclopædia Britannica 2006 Ultimate Reference Suite DVD 2009 In many vertebrates, the sclera is reinforced with plates of
cartilage Cartilage is a resilient and smooth type of connective tissue. In tetrapods, it covers and protects the ends of long bones at the joints as articular cartilage, and is a structural component of many body parts including the rib cage, the neck an ...
or bone, together forming a circular structure called the sclerotic ring. In primitive fish, this ring consists of four plates, but the number is lower in many living ray-finned fishes, and much higher in lobe-finned fishes, various
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 birds. The ring has disappeared in many groups, including living
amphibian Amphibians are tetrapod, four-limbed and ectothermic vertebrates of the Class (biology), class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terres ...
s, some reptiles and fish, and all
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
s. The eyes of all non-human primates are dark with small, barely visible sclera.


Histology

The
collagen Collagen () is the main structural protein in the extracellular matrix found in the body's various connective tissues. As the main component of connective tissue, it is the most abundant protein in mammals, making up from 25% to 35% of the whole ...
of the sclera is continuous with the cornea. From outer to innermost, the four layers of the sclera are: * episclera * stroma * lamina fusca * endothelium The sclera is opaque due to the irregularity of the Type I collagen fibers, as opposed to the near-uniform thickness and parallel arrangement of the corneal collagen. Moreover, the cornea bears more mucopolysaccharide (a carbohydrate that has among its repeating units a nitrogenous sugar, hexosamine) to embed the fibrils. The cornea, unlike the sclera, has five layers. The middle, thickest layer is also called the stroma. The sclera, like the cornea, contains a basal endothelium, above which there is the lamina fusca, containing a high count of pigment cells. Sometimes, very small gray-blue spots can appear on the sclera, a harmless condition called scleral melanocytosis.


Function

Human eyes are somewhat distinctive in the animal kingdom in that the sclera is very plainly visible whenever the eye is open. This is not just due to the white color of the human sclera, which many other species share, but also to the fact that the human iris is relatively small and comprises a significantly smaller portion of the exposed eye surface compared to other animals. It is theorized that this adaptation evolved because of our social nature as the eye became a useful communication tool in addition to a
sensory organ A sense is a biological system used by an organism for sensation, the process of gathering information about the world through the detection of stimuli. (For example, in the human body, the brain which is part of the central nervous system rec ...
. It is believed that the conspicuous sclera of the human eye makes it easier for one individual to identify where another individual is looking, increasing the efficacy of this particular form of nonverbal communication, called
cooperative eye hypothesis The cooperative eye hypothesis is a proposed explanation for the appearance of the human eye. It suggests that the eye's distinctive visible characteristics evolved to make it easier for humans to follow another's gaze while communicating or while ...
.Michael Tomasello, Brian Hare, Hagen Lehmann, Josep Call. "Reliance on head versus eyes in the gaze following of great apes and human infants: the cooperative eye hypothesis" http://www.chrisknight.co.uk/wp-content/uploads/2008/06/eyes-cooperation.pdf Animal researchers have also found that, in the course of their domestication, dogs have also developed the ability to pick up visual cues from the eyes of humans. Dogs do not seem to use this form of communication with one another and only look for visual information from the eyes of humans.


Injury


Trauma

The bony area that makes up the human eye socket provides exceptional protection to the sclera. However, if the sclera is ruptured by a blunt force or is penetrated by a sharp object, the recovery of full former vision is usually rare. If pressure is applied slowly, the eye is actually very elastic. However, most ruptures involve objects moving at some velocity. The cushion of orbital fat protects the sclera from head-on blunt forces, but damage from oblique forces striking the eye from the side is not prevented by this cushion. Hemorrhaging and a dramatic drop in intraocular pressure are common, along with a reduction in visual perception to only broad hand movements and the presence or absence of light. However, a low-velocity injury which does not puncture and penetrate the sclera requires only superficial treatment and the removal of the object. Sufficiently small objects which become embedded and which are subsequently left untreated may eventually become surrounded by a benign cyst, causing no other damage or discomfort.


Thermal trauma

The sclera is rarely damaged by brief exposure to heat: the eyelids provide exceptional protection, and the fact that the sclera is covered in layers of moist tissue means that these tissues are able to cause much of the offending heat to become dissipated as steam before the sclera itself is damaged. Even relatively low-temperature molten metals when splashed against an open eye have been shown to cause very little damage to the sclera, even while creating detailed casts of the surrounding eyelashes. Prolonged exposure, however—on the order of 30 seconds—at temperatures above will begin to cause scarring, and above will cause extreme changes in the sclera and surrounding tissue. Such long exposures even in industrial settings are virtually nonexistent.


Chemical injury

The sclera is highly resistant to injury from brief exposure to toxic chemicals. The reflexive production of tears at the onset of chemical exposure tends to quickly wash away such irritants, preventing further harm. Acids with a pH below 2.5 are the source of greatest acidic burn risk, with
sulfuric acid Sulfuric acid (American spelling and the preferred IUPAC name) or sulphuric acid ( Commonwealth spelling), known in antiquity as oil of vitriol, is a mineral acid composed of the elements sulfur, oxygen and hydrogen, with the molecular formu ...
, the kind present in car batteries and therefore commonly available, being among the most dangerous in this regard. However, acid burns, even severe ones, seldom result in loss of the eye. Alkali burns, on the other hand, such as those resulting from exposure to
ammonium hydroxide Ammonia solution, also known as ammonia water, ammonium hydroxide, ammoniacal liquor, ammonia liquor, aqua ammonia, aqueous ammonia, or (inaccurately) ammonia, is a solution of ammonia in water. It can be denoted by the symbols NH3(aq). Although ...
or ammonium chloride or other chemicals with a pH above 11.5, will cause cellular tissue in the sclera to
saponify Saponification is a process of converting esters into soaps and alcohols by the action of aqueous alkali (for example, aqueous sodium hydroxide solutions). Soaps are salts of fatty acids, which in turn are carboxylic acids with long carbon chains. ...
and should be considered medical emergencies requiring immediate treatment.


Abnormal coloring

Redness of the sclera is typically caused by
eye irritation The human eye is a sensory organ, part of the sensory nervous system, that reacts to visible light and allows humans to use visual information for various purposes including seeing things, keeping balance, and maintaining circadian rhythm. T ...
causing blood vessels to expand, such as in conjunctivitis ("pink eye"). Episcleritis is a generally benign condition of the episclera causing eye redness.
Scleritis Scleritis is a serious inflammatory disease that affects the white outer coating of the eye, known as the sclera. The disease is often contracted through association with other diseases of the body, such as granulomatosis with polyangiitis or rhe ...
is a serious inflammatory disease of the sclera causing redness of the sclera often progressing to purple. Yellowing or a light green color of the sclera is a visual symptom of
jaundice Jaundice, also known as icterus, is a yellowish or greenish pigmentation of the skin and sclera due to high bilirubin levels. Jaundice in adults is typically a sign indicating the presence of underlying diseases involving abnormal heme meta ...
. In cases of osteogenesis imperfecta, the sclera may appear to have a blue tint, more pronounced than the slight blue tint seen in children. The blue tint is caused by the showing of the underlying uveal tract (
choroid The choroid, also known as the choroidea or choroid coat, is a part of the uvea, the vascular layer of the eye, and contains connective tissues, and lies between the retina and the sclera. The human choroid is thickest at the far extreme rear ...
and retinal pigment epithelium). In very rare but severe cases of kidney failure and liver failure, the sclera may turn black.


See also

* Extraocular implant *
Scleral tattooing Scleral tattooing is the practice of tattooing the sclera, or white part of the human eye. The dye is not injected into the tissue, but between two layers of the eye, where it spreads out over a large area. The process is not common as there a ...
*
Sclerotomy Sclerotomy is a medical intervention that involves surgical cutting in the white area of the eye, known as the sclera. The goal of this intervention is usually done to correct defects in sclera that resulted as a complication of glaucoma of other ...


Notes


References


External links

* * —"Sagittal Section Through the Eyeball" * {{Authority control Eye Human eye anatomy