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. 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, 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 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 choroidal tissue to form a plate ( lamina cribrosa) across the optic nerve with perforations through which the optic fibers ( fasciculi) 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 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 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. 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 fishes, 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

. From outer to innermost, the four layers of the sclera are: *

* 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 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

, 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, 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 eyelashe