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
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 ...
containing mainly
collagen
Collagen () is the main structural protein in the extracellular matrix of the connective tissues of many animals. It is the most abundant protein in mammals, making up 25% to 35% of protein content. Amino acids are bound together to form a trip ...
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 the development of the
embryo
An embryo ( ) is the initial stage of development for 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 sp ...
, the sclera is derived from the
neural crest
The neural crest is a ridge-like structure that is formed transiently between the epidermal ectoderm and neural plate during vertebrate development. Neural crest cells originate from this structure through the epithelial-mesenchymal transition, ...
. 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 (; ) is a family of biomolecules organized as oligomers or polymers, which among other functions provide the pigments of many organisms. Melanin pigments are produced in a specialized group of cells known as melanocytes.
There are ...
pigmentation.
In
human
Humans (''Homo sapiens'') or modern humans are the most common and widespread species of primate, and the last surviving species of the genus ''Homo''. They are Hominidae, great apes characterized by their Prehistory of nakedness and clothing ...
s, and some other
vertebrates
Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain.
The vertebrates make up the subphylum Vertebra ...
, the whole sclera is white or pale, contrasting with the coloured
iris. The
cooperative eye hypothesis suggests that the pale sclera evolved as a method of
nonverbal communication
Nonverbal communication is the transmission of messages or signals through a nonverbal platform such as eye contact (oculesics), body language (kinesics), social distance (proxemics), touch (Haptic communication, haptics), voice (prosody (lingui ...
that makes it easier for one individual to identify where another individual is looking. Other mammals with white or pale sclera include
chimpanzee
The chimpanzee (; ''Pan troglodytes''), also simply known as the chimp, is a species of Hominidae, great ape native to the forests and savannahs of tropical Africa. It has four confirmed subspecies and a fifth proposed one. When its close rel ...
s, many
orangutan
Orangutans are great apes native to the rainforests of Indonesia and Malaysia. They are now found only in parts of Borneo and Sumatra, but during the Pleistocene they ranged throughout Southeast Asia and South China. Classified in the genus ...
s, some
gorilla
Gorillas are primarily herbivorous, terrestrial great apes that inhabit the tropical forests of equatorial Africa. The genus ''Gorilla'' is divided into two species: the eastern gorilla and the western gorilla, and either four or five su ...
s, and
bonobo
The bonobo (; ''Pan paniscus''), also historically called the pygmy chimpanzee (less often the dwarf chimpanzee or gracile chimpanzee), is an endangered great ape and one of the two species making up the genus ''Pan (genus), Pan'' (the other bei ...
s.
Structure
The sclera forms the posterior five-sixths of the
connective tissue
Connective tissue is one of the four primary types of animal tissue, a group of cells that are similar in structure, along with epithelial tissue, muscle tissue, and nervous tissue. It develops mostly from the mesenchyme, derived from the mesod ...
coat of the
human eyeball. It is continuous with the
dura mater and the
cornea
The cornea is the transparency (optics), transparent front part of the eyeball which covers the Iris (anatomy), iris, pupil, and Anterior chamber of eyeball, anterior chamber. Along with the anterior chamber and Lens (anatomy), lens, the cornea ...
, and maintains the shape of the eyeball, 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
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 system, visual information from the retina to the brain. In humans, the optic nerve i ...
. 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 t ...
, 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. It contains connective tissues, and lies between the retina and the sclera. The human choroid is thickest at the far extreme rear o ...
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 to 0.3 mm just behind the insertions of the
four rectus muscles. The sclera's blood vessels are mainly on the surface. Along with the vessels of the
conjunctiva
In the anatomy of the eye, the conjunctiva (: conjunctivae) 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 gobl ...
(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
An encyclopedia is a reference work or compendium providing summaries of knowledge, either general or special, in a particular field or discipline. Encyclopedias are divided into articles or entries that are arranged alphabetically by artic ...
2009
In many
vertebrate
Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain.
The vertebrates make up the subphylum Vertebra ...
s, the sclera is reinforced with plates of
cartilage
Cartilage is a resilient and smooth type of connective tissue. Semi-transparent and non-porous, it is usually covered by a tough and fibrous membrane called perichondrium. In tetrapods, it covers and protects the ends of long bones at the joints ...
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, ...
, together forming a circular structure called the
sclerotic ring
The scleral ring or sclerotic ring is a hardened ring of plates, often derived from bone, that is found in the eyes of many animals in several groups of vertebrates. Some species of mammals, amphibians, and crocodilians lack scleral rings. The rin ...
. 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 fish or actinopterygians, is a class of bony fish that comprise over 50% of living vertebrate species. They are so called because of their lightly built fins made of webbings of sk ...
es, and much higher in
lobe-finned fish
Sarcopterygii (; )—sometimes considered synonymous with Crossopterygii ()—is a clade (traditionally a class or subclass) of vertebrate animals which includes a group of bony fish commonly referred to as lobe-finned fish. These vertebrates ar ...
es,
crocodilians
Crocodilia () is an order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorph pseudosuchi ...
, various
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, and
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. The ring has disappeared in many groups, including living
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, some reptiles and fish, and all
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.
The eyes of all non-human
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 have been thought to be dark with small, barely visible sclera, but recent research has suggested that white sclera are not uncommon in chimpanzees, and are also present in other mammals.
Histology
Histologically speaking, it is characterized as
dense connective tissue
Dense connective tissue, also called dense fibrous tissue, is a type of connective tissue with fibers as its main extracellular matrix, matrix element. The fibers are mainly composed of type I collagen. Crowded between the collagen fibers are rows ...
made primarily of
type 1 collagen fibers. The
collagen
Collagen () is the main structural protein in the extracellular matrix of the connective tissues of many animals. It is the most abundant protein in mammals, making up 25% to 35% of protein content. Amino acids are bound together to form a trip ...
of the sclera is continuous with the
cornea
The cornea is the transparency (optics), transparent front part of the eyeball which covers the Iris (anatomy), iris, pupil, and Anterior chamber of eyeball, anterior chamber. Along with the anterior chamber and Lens (anatomy), lens, the cornea ...
. From outer to innermost, the four layers of the sclera are:
*
episclera
*
stroma
*
lamina fusca
*
endothelium
The endothelium (: endothelia) 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 r ...
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 six layers. The middle (third in number; provided the first layer being the anterior and outermost and the sixth layer being the posterior and the inner most), 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 Biology, biological Kingdom (biology), kingdom Animalia (). With few exceptions, animals heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, ...
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 humans’
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 fro ...
nature as the eye became a useful
communication
Communication is commonly defined as the transmission of information. Its precise definition is disputed and there are disagreements about whether Intention, unintentional or failed transmissions are included and whether communication not onl ...
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 surroundings through the detection of stimuli. Although, in some cultures, five human senses were traditionally identified as su ...
. It is believed that the exposed 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 is the transmission of messages or signals through a nonverbal platform such as eye contact (oculesics), body language (kinesics), social distance (proxemics), touch (Haptic communication, haptics), voice (prosody (lingui ...
, called
cooperative eye hypothesis. 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.
Another hypothesis states that the distinctiveness of the human's sclera is due to
genetic drift
Genetic drift, also known as random genetic drift, allelic drift or the Wright effect, is the change in the Allele frequency, frequency of an existing gene variant (allele) in a population due to random chance.
Genetic drift may cause gene va ...
and
sexual selection
Sexual selection is a mechanism of evolution in which members of one sex mate choice, choose mates of the other sex to mating, mate with (intersexual selection), and compete with members of the same sex for access to members of the opposite sex ...
. A visibly white sclera is perceived as a sign of good health and youthfullness, this could have been a criterion of selection when looking for a mate. In this case, humans' ability to communicate with their eyes (glancing, cluing at others) would only be a consequence of a very visible sclera.
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 bubb ...
, 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 (English in the Commonwealth of Nations, Commonwealth spelling), known in antiquity as oil of vitriol, is a mineral acid composed of the elements sulfur, oxygen, ...
, 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
Ammonium chloride is an inorganic chemical compound with the chemical formula , also written as . It is an ammonium salt of hydrogen chloride. It consists of ammonium cations and chloride anions . It is a white crystalline salt (chemistry), sal ...
or other chemicals with a pH above 11.5, will cause cellular tissue in the sclera to
saponify and should be considered medical emergencies requiring immediate treatment.
Abnormal coloring
Redness of the sclera is typically caused by
eye irritation causing blood vessels to expand, such as in
conjunctivitis
Conjunctivitis, also known as pink eye or Madras eye, is inflammation of the conjunctiva, the thin, clear layer that covers the white surface of the eye and the inner eyelid. It makes the eye appear pink or reddish. Pain, burning, scratchiness ...
("pink eye").
Episcleritis is a generally benign condition of the
episclera causing eye redness.
Scleritis 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, less frequently, 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 ...
.
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 bone fracture, break easily. The range of symptoms—on the skeleton as well as on the body's other Or ...
, 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. It contains connective tissues, and lies between the retina and the sclera. The human choroid is thickest at the far extreme rear o ...
and
retinal pigment epithelium
The pigmented layer of retina or retinal pigment epithelium (RPE) is the pigment
A pigment is a powder used to add or alter color or change visual appearance. Pigments are completely or nearly solubility, insoluble and reactivity (chemistry), ...
).
In those with
Ehlers-Danlos syndrome, the sclera may be tinted blue due to the lack of proper connective tissue.
[Asanad, S., Bayomi, M., Brown, D., Buzzard, J., Lai, E., Ling, C., Miglani, T., Mohammed, T., Tsai, J., Uddin, O., & Singman, E. (2022). Ehlers-Danlos syndromes and their manifestations in the visual system. Frontiers in Medicine, 9. https://doi.org/10.3389/fmed.2022.996458]
In very rare but severe cases of
kidney failure
Kidney failure, also known as renal failure or end-stage renal disease (ESRD), 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 fa ...
and
liver failure
Liver failure is the inability of the liver to perform its normal synthetic and metabolic functions as part of normal physiology. Two forms are recognised, acute and chronic (cirrhosis). Recently, a third form of liver failure known as acute- ...
, the sclera may turn black.
See also
*
Extraocular implant
*
Extraocular muscles
The extraocular muscles, or extrinsic ocular muscles, are the seven extrinsic muscles of the eye in human eye, humans and other animals. Six of the extraocular muscles, the four recti muscles, and the superior oblique muscle, superior and inferior ...
** Rectus muscles:
***
Inferior rectus muscle
The inferior rectus muscle is a muscle in the orbit near the eye. It is one of the four recti muscles in the group of extraocular muscles. It originates from the common tendinous ring, and inserts into the anteroinferior surface of the eye. It d ...
***
Lateral rectus muscle
The lateral rectus muscle is a muscle on the lateral side of the eye in the orbit. It is one of six extraocular muscles that control the movements of the eye. The lateral rectus muscle is responsible for lateral movement of the eyeball, specif ...
***
Medial rectus muscle
The medial rectus muscle is a muscle in the orbit near the eye. It is one of the extraocular muscles. It originates from the common tendinous ring, and inserts into the anteromedial surface of the eye. It is supplied by the inferior division of t ...
***
Superior rectus muscle
The superior rectus muscle is a muscle in the orbit. It is one of the extraocular muscles. It is innervated by the superior division of the oculomotor nerve (III). In the primary position (looking straight ahead), its primary function is elevatio ...
*
Scleral tattooing
*
Sclerotomy
Notes
References
External links
*
* —"Sagittal Section Through the Eyeball"
*
{{Authority control
Eye
Human eye anatomy