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 containing mainly
collagen 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 ai ...
. In humans, and some other
vertebrates
Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, wi ...
, the whole sclera is white, contrasting with the coloured
iris
Iris most often refers to:
*Iris (anatomy), part of the eye
*Iris (mythology), a Greek goddess
* ''Iris'' (plant), a genus of flowering plants
* Iris (color), an ambiguous color term
Iris or IRIS may also refer to:
Arts and media
Fictional ent ...
, but in most
mammals
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), fu ...
, 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 sperm ...
, the sclera is derived from the
neural crest
Neural crest cells are a temporary group of cells unique to vertebrates that arise from the embryonic ectoderm germ layer, and in turn give rise to a diverse cell lineage—including melanocytes, craniofacial cartilage and bone, smooth muscle, ...
. 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
Melanin (; from el, μέλας, melas, black, dark) is a broad term for a group of natural pigments found in most organisms. Eumelanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the amin ...
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 suggests this has evolved as a method of
nonverbal communication
Nonverbal communication (NVC) is the transmission of messages or signals through a nonverbal platform such as eye contact, facial expressions, gestures, posture, and body language. It includes the use of social cues, kinesics, distance (p ...
.
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 tissue ...
coat of the
globe
A globe is a spherical model of Earth, of some other celestial body, or of the celestial sphere. Globes serve purposes similar to maps, but unlike maps, they do not distort the surface that they portray except to scale it down. A model glo ...
. It is continuous with the
dura mater
In neuroanatomy, dura mater is a thick membrane made of dense irregular connective tissue that surrounds the brain and spinal cord. It is the outermost of the three layers of membrane called the meninges that protect the central nervous syste ...
and the
cornea
The cornea is the transparent front part of the eye that covers the iris, pupil, and anterior chamber. Along with the anterior chamber and lens, the cornea refracts light, accounting for approximately two-thirds of the eye's total optical ...
, and maintains the shape of the globe, offering resistance to internal and external forces, and provides an attachment for the
extraocular muscle
The extraocular muscles (extrinsic ocular muscles), are the seven extrinsic muscles of the human eye. Six of the extraocular muscles, the four recti muscles, and the superior and inferior oblique muscles, control movement of the eye and the o ...
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
In neuroanatomy, the optic nerve, also known as the second cranial nerve, cranial nerve II, or simply CN II, is a paired cranial nerve that transmits visual information from the retina to the brain. In humans, the optic nerve is derived fro ...
. 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 th ...
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 fasc ...
) pass. The thickness of the sclera varies from 1 mm at the
posterior pole In ophthalmology, the posterior pole is the back of the eye, usually referring to the retina between the optic disc and the macula.Cassin, B. and Solomon, S. ''Dictionary of Eye Terminology''. Gainesville, Florida: Triad Publishing Company, 1990.
S ...
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
The conjunctiva is a thin mucous membrane that lines the inside of the eyelids and covers the sclera (the white of the eye). It is composed of non-keratinized, stratified squamous epithelium with goblet cells, stratified columnar epitheli ...
(which is a thin layer covering the sclera), those in the
episclera
The episclera is the outermost layer of the sclera (the white of the eye).Cassin, B. and Solomon, S. ''Dictionary of Eye Terminology''. Gainesville, Florida: Triad Publishing Company, 1990. It is composed of loose, fibrous, elastic tissue and atta ...
render the inflamed eye bright red.
["eye, human."Encyclopædia Britannica from ]Encyclopædia Britannica 2006 Ultimate Reference Suite DVD
An encyclopedia (American English) or encyclopædia (British English) is a reference work or compendium providing summaries of knowledge either general or special to a particular field or discipline. Encyclopedias are divided into articles ...
2009
In many
vertebrate
Vertebrates () comprise all animal taxon, taxa within the subphylum Vertebrata () (chordates with vertebral column, backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the ...
s, the sclera is reinforced with plates of
cartilage or
bone
A bone is a rigid organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red and white blood cells, store minerals, provide structure and support for the body, an ...
, together forming a circular structure called the
sclerotic ring
Sclerotic rings are rings of bone found in the eyes of many animals in several groups of vertebrates, except for mammals and crocodilians. They can be made up of single bones or multiple segments and take their name from the sclera. They are bel ...
. In primitive fish, this ring consists of four plates, but the number is lower in many living
ray-finned fish
Actinopterygii (; ), members of which are known as ray-finned fishes, is a class of bony fish. They comprise over 50% of living vertebrate species.
The ray-finned fishes are so called because their fins are webs of skin supported by bony or h ...
es, and much higher in
lobe-finned fish
Sarcopterygii (; ) — sometimes considered synonymous with Crossopterygii () — is a taxon (traditionally a class or subclass) of the bony fishes known as the lobe-finned fishes. The group Tetrapoda, a mostly terrestrial superclass includi ...
es, various
reptiles, and
bird
Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
s. The ring has disappeared in many groups, including living
amphibians, some reptiles and fish, and all
mammals.
The eyes of all non-human
primate
Primates are a diverse order (biology), order of mammals. They are divided into the Strepsirrhini, strepsirrhines, which include the lemurs, galagos, and lorisids, and the Haplorhini, haplorhines, which include the Tarsiiformes, tarsiers and ...
s are dark with small, barely visible sclera.
Histology
The
collagen of the sclera is continuous with the
cornea
The cornea is the transparent front part of the eye that covers the iris, pupil, and anterior chamber. Along with the anterior chamber and lens, the cornea refracts light, accounting for approximately two-thirds of the eye's total optical ...
. From outer to innermost, the four layers of the sclera are:
*
episclera
The episclera is the outermost layer of the sclera (the white of the eye).Cassin, B. and Solomon, S. ''Dictionary of Eye Terminology''. Gainesville, Florida: Triad Publishing Company, 1990. It is composed of loose, fibrous, elastic tissue and atta ...
*
stroma
*
lamina fusca
*
endothelium
The endothelium is a single layer of squamous endothelial cells that line the interior surface of blood vessels and lymphatic vessels. The endothelium forms an interface between circulating blood or lymph in the lumen and the rest of the ve ...
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
Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...
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
Iris most often refers to:
*Iris (anatomy), part of the eye
*Iris (mythology), a Greek goddess
* ''Iris'' (plant), a genus of flowering plants
* Iris (color), an ambiguous color term
Iris or IRIS may also refer to:
Arts and media
Fictional ent ...
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
Social organisms, including human(s), live collectively in interacting populations. This interaction is considered social whether they are aware of it or not, and whether the exchange is voluntary or not.
Etymology
The word "social" derives from ...
nature as the eye became a useful
communication
Communication (from la, communicare, meaning "to share" or "to be in relation with") is usually defined as the transmission of information. The term may also refer to the message communicated through such transmissions or the field of inqu ...
tool in addition to a
sensory organ. 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
Nonverbal communication (NVC) is the transmission of messages or signals through a nonverbal platform such as eye contact, facial expressions, gestures, posture, and body language. It includes the use of social cues, kinesics, distance (p ...
, called
cooperative eye hypothesis.
[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
A cyst is a closed sac, having a distinct envelope and division compared with the nearby tissue. Hence, it is a cluster of cells that have grouped together to form a sac (like the manner in which water molecules group together to form a bubble ...
, 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, 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.
...
causing blood vessels to expand, such as in
conjunctivitis
Conjunctivitis, also known as pink eye, is inflammation of the outermost layer of the white part of the eye and the inner surface of the eyelid. It makes the eye appear pink or reddish. Pain, burning, scratchiness, or itchiness may occur. The ...
("pink eye").
Episcleritis
Episcleritis is a benign, self-limiting inflammatory disease affecting part of the eye called the episclera. The episclera is a thin layer of tissue that lies between the conjunctiva and the connective tissue layer that forms the white of the ey ...
is a generally benign condition of the
episclera
The episclera is the outermost layer of the sclera (the white of the eye).Cassin, B. and Solomon, S. ''Dictionary of Eye Terminology''. Gainesville, Florida: Triad Publishing Company, 1990. It is composed of loose, fibrous, elastic tissue and atta ...
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 met ...
.
In cases of
osteogenesis imperfecta
Osteogenesis imperfecta (; OI), colloquially known as brittle bone disease, is a group of genetic disorders that all result in bones that break easily. The range of symptoms—on the skeleton as well as on the body's other organs—may b ...
, 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
The uvea (; Latin, Lat. ''uva'', "grape"), also called the ''uveal layer'', ''uveal coat'', ''uveal tract'', ''vascular tunic'' or ''vascular layer'' is the pigmented middle of the three concentric layers that make up an human eye, eye.
Hist ...
(
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
The pigmented layer of retina or retinal pigment epithelium (RPE) is the pigmented cell layer just outside the neurosensory retina that nourishes retinal visual cells, and is firmly attached to the underlying choroid and overlying retinal visual ...
).
In very rare but severe cases of
kidney failure
Kidney failure, also known as end-stage kidney disease, is a medical condition in which the kidneys can no longer adequately filter waste products from the blood, functioning at less than 15% of normal levels. Kidney failure is classified as eit ...
and
liver failure, the sclera may turn black.
See also
*
Extraocular implant
An extraocular implant (also known as eyeball jewelry) is a cosmetic implant involving a tiny piece of decorative jewelry which is implanted within the superficial, interpalpebral conjunctiva or sclera of the human eye.
History and culture
Eyeb ...
*
Scleral tattooing
*
Sclerotomy
Notes
References
External links
*
* —"Sagittal Section Through the Eyeball"
*
{{Authority control
Eye
Human eye anatomy