The human eye
1. vitreous body 2. ora serrata 3. ciliary muscle 4. ciliary zonules 5. canal of Schlemm 6. pupil 7. anterior chamber 8. cornea 9. iris 10. lens cortex 11. lens nucleus 12. ciliary process 13. conjunctiva 14. inferior oblique muscle 15. inferior rectus muscle 16. medial rectus muscle 17. retinal arteries and veins 18. optic disc 19. dura mater 20. central retinal artery 21. central retinal vein 22. optic nerve 23. vorticose vein 24. bulbar sheath 25. macula 26. fovea 27. sclera 28. choroid 29. superior rectus muscle 30. retina

The human eye is an organ which reacts to light for several purposes. As a conscious sense organ, the mammalian eye allows vision. Rod and cone cells in the retina allow conscious light perception and vision including color differentiation and the perception of depth. The human eye can distinguish about 10 million colors.^[1]

In common with the eyes of other mammals, the human eye's non-image-forming photosensitive ganglion cells in the retina receive the light signals which affect adjustment of the size of the pupil, regulation and suppression of the hormone melatonin and entrainment of the body clock.^[2]

General properties[link]

A human eye.

The eye is not properly a sphere, rather it is a fused two-piece unit. The smaller frontal unit, more curved, called the cornea is linked to the larger unit called the sclera. The corneal segment is typically about 8 mm (0.3 in) in radius. The sclerotic chamber constitutes the remaining five-sixths; its radius is typically about 12 mm. The cornea and sclera are connected by a ring called the limbus. The iris – the color of the eye – and its black center, the pupil, are seen instead of the cornea due to the cornea's transparency. To see inside the eye, an ophthalmoscope is needed, since light is not reflected out. The fundus (area opposite the pupil) shows the characteristic pale optic disk (papilla), where vessels entering the eye pass across and optic nerve fibers depart the globe.

Dimensions[link]

The dimensions differ among adults by only one or two millimeters. The vertical measure, generally less than the horizontal distance, is about 24 mm among adults, at birth about 16–17 mm. (about 0.65 inch) The eyeball grows rapidly, increasing to 22.5–23 mm (approx. 0.89 in) by the age of three years. From then to age 13, the eye attains its full size. The volume is 6.5 ml (0.4 cu. in.) and the weight is 7.5 g (0.25 oz.).^{[citation needed]}

Components[link]

The eye is made up of three coats, enclosing three transparent structures. The outermost layer is composed of the cornea and sclera. The middle layer consists of the choroid, ciliary body, and iris. The innermost is the retina, which gets its circulation from the vessels of the choroid as well as the retinal vessels, which can be seen in an ophthalmoscope.

Within these coats are the aqueous humor, the vitreous body, and the flexible lens. The aqueous humor is a clear fluid that is contained in two areas: the anterior chamber between the cornea and the iris and exposed area of the lens. The lens is suspended to the ciliary body by the suspensory ligament (Zonule of Zinn), made up of fine transparent fibers. The vitreous body, the posterior chamber, is a clear jelly that is much larger than the aqueous humor, present behind lens and the rest, and is bordered by the sclera, zonule, and lens. They are connected via the pupil.^[3]

Dynamic range[link]

The retina has a static contrast ratio of around 100:1 (about 6.5 f-stops). As soon as the eye moves (saccades) it re-adjusts its exposure both chemically and geometrically by adjusting the iris which regulates the size of the pupil. Initial dark adaptation takes place in approximately four seconds of profound, uninterrupted darkness; full adaptation through adjustments in retinal chemistry (the Purkinje effect) are mostly complete in thirty minutes. Hence, a dynamic contrast ratio of about 1,000,000:1 (about 20 f-stops) is possible.^[4] The process is nonlinear and multifaceted, so an interruption by light merely starts the adaptation process over again. Full adaptation is dependent on good blood flow; thus dark adaptation may be hampered by poor circulation, and vasoconstrictors like tobacco.^{[citation needed]}

The eye includes a lens not dissimilar to lenses found in optical instruments such as cameras and the same principles can be applied. The pupil of the human eye is its aperture; the iris is the diaphragm that serves as the aperture stop. Refraction in the cornea causes the effective aperture (the entrance pupil) to differ slightly from the physical pupil diameter. The entrance pupil is typically about 4 mm in diameter, although it can range from 2 mm (f/8.3) in a brightly lit place to 8 mm (f/2.1) in the dark. The latter value decreases slowly with age, older people's eyes sometimes dilate to not more than 5-6mm.^{[citation needed]}

Field of view[link]

The approximate field of view of a human eye is 95° out, 75° down, 60° in, 60° up.^{[clarification needed]} About 12–15° temporal and 1.5° below the horizontal is the optic nerve or blind spot which is roughly 7.5° high and 5.5° wide.^[5]

Eye irritation[link]

Bloodshot eyeball

Eye irritation has been defined as “the magnitude of any stinging, scratching, burning, or other irritating sensation from the eye”.^[6] It is a common problem experienced by people of all ages. Related eye symptoms and signs of irritation are e.g. discomfort, dryness, excess tearing, itching, grating, sandy sensation, smarting, ocular fatigue, pain, scratchiness, soreness, redness, swollen eyelids, and tiredness, etc. These eye symptoms are reported with intensities from severe to less severe. It has been suggested that these eye symptoms are related to different causal mechanisms.^[7]

Several suspected causal factors in our environment have been studied so far.^[8] One hypothesis is that indoor air pollution may cause eye and airway irritation.^[9][10] Eye irritation depends somewhat on destabilization of the outer-eye tear film, in which the formation of dry spots results in such ocular discomfort as dryness.^[9][11][12] Occupational factors are also likely to influence the perception of eye irritation. Some of these are lighting (glare and poor contrast), gaze position, a limited number of breaks, and a constant function of accommodation, musculoskeletal burden, and impairment of the visual nervous system.^[13][14] Another factor that may be related is work stress.^[15][16] In addition, psychological factors have been found in multivariate analyses to be associated with an increase in eye irritation among VDU users.^[17][18] Other risk factors, such as chemical toxins/irritants, e.g. amines, formaldehyde, acetaldehyde, acrolein, N-decane, VOCs; ozone, pesticides and preservatives, allergens, etc. might cause eye irritation as well.

Certain volatile organic compounds that are both chemically reactive and airway irritants may cause eye irritation as well. Personal factors (e.g., use of contact lenses, eye make-up, and certain medications) may also affect destabilization of the tear film and possibly result in more eye symptoms.^[7] Nevertheless, if airborne particles alone should destabilize the tear film and cause eye irritation, their content of surface-active compounds must be high.^[7] An integrated physiological risk model with blink frequency, destabilization, and break-up of the eye tear film as inseparable phenomena may explain eye irritation among office workers in terms of occupational, climate, and eye-related physiological risk factors.^[7]

There are two major measures of eye irritation. One is blink frequency which can be observed by human behavior. The other measures are break up time, tear flow, hyperemia (redness, swelling), tear fluid cytology, and epithelial damage (vital stains) etc., which are human beings’ physiological reactions. Blink frequency is defined as the number of blinks per minute and it is associated with eye irritation. Blink frequencies are individual with mean frequencies of < 2-3 to 20-30 blinks/minute, and they depend on environmental factors including the use of contact lenses. Dehydration, mental activities, work conditions, room temperature, relative humidity, and illumination all influence blink frequency. Break-up time (BUT) is another major measure of eye irritation and tear film stability.^[19] It is defined as the time interval (in seconds) between blinking and rupture. BUT is considered to reflect the stability of the tear film as well. In normal persons, the break-up time exceeds the interval between blinks, and, therefore, the tear film is maintained.^[20] Studies have shown that blink frequency is correlated negatively with break-up time. This phenomenon indicates that perceived eye irritation is associated with an increase in blink frequency since the cornea and conjunctiva both have sensitive nerve endings that belong to the first trigeminal branch.^[21][22] Other evaluating methods, such as hyperemia, cytology etc. have increasingly been used to assess eye irritation.

There are other factors that related to eye irritation as well. Three major factors that influence the most are indoor air pollution, contact lenses and gender differences. Field studies have found that the prevalence of objective eye signs is often significantly altered among office workers in comparisons with random samples of the general population.^{[23][24][25][26]} These research results might indicate that indoor air pollution has played an important role in causing eye irritation. There are more and more people wearing contact lens now and dry eyes appear to be the most common complaint among contact lens wearers.^[27][28][29] Although both contact lens wearers and spectacle wearers experience similar eye irritation symptoms, dryness, redness, and grittiness have been reported far more frequently among contact lens wearers and with greater severity than among spectacle wearers.^[29] Studies have shown that incidence of dry eyes increases with age.^[30][31] especially among women.^[32] Tear film stability (e.g. break-up time) is significantly lower among women than among men. In addition, women have a higher blink frequency while reading.^[33] Several factors may contribute to gender differences. One is the use of eye make-up. Another reason could be that the women in the reported studies have done more VDU work than the men, including lower grade work. A third often-quoted explanation is related to the age-dependent decrease of tear secretion, particularly among women after 40 years of age.,^[34][35][36]

In a study conducted by UCLA, the frequency of reported symptoms in industrial buildings was investigated.^[37] The study's results were that eye irritation was the most frequent symptom in industrial building spaces, at 81%. Modern office work with use of office equipment has raised concerns about possible adverse health effects.^[38] Since the 1970s, reports have linked mucosal, skin, and general symptoms to work with self-copying paper. Emission of various particulate and volatile substances has been suggested as specific causes. These symptoms have been related to Sick Building Syndrome (SBS), which involves symptoms such as irritation to the eyes, skin, and upper airways, headache and fatigue.^[39]

Many of the symptoms described in SBS and multiple chemical sensitivity (MCS) resemble the symptoms known to be elicited by airborne irritant chemicals.^[40] A repeated measurement design was employed in the study of acute symptoms of eye and respiratory tract irritation resulting from occupational exposure to sodium borate dusts.^[41] The symptom assessment of the 79 exposed and 27 unexposed subjects comprised interviews before the shift began and then at regular hourly intervals for the next six hours of the shift, four days in a row.^[41] Exposures were monitored concurrently with a personal real time aerosol monitor. Two different exposure profiles, a daily average and short term (15 minute) average, were used in the analysis. Exposure-response relations were evaluated by linking incidence rates for each symptom with categories of exposure.^[41]

Acute incidence rates for nasal, eye, and throat irritation, and coughing and breathlessness were found to be associated with increased exposure levels of both exposure indices. Steeper exposure-response slopes were seen when short term exposure concentrations were used. Results from multivariate logistic regression analysis suggest that current smokers tended to be less sensitive to the exposure to airborne sodium borate dust.^[41]

Several actions can be taken to prevent eye irritation—

• trying to maintain normal blinking by avoiding room temperatures that are too high; avoiding relative humidities that are too high or too low, because they reduce blink frequency or may increase water evaporation^[7]
• trying to maintain an intact tear film by the following actions. 1) blinking and short breaks may be beneficial for VDU users.^[42][43] Increase these two actions might help maintain the tear film. 2) downward gazing is recommended to reduce the ocular surface area and water evaporation.^[44][45][46] 3) the distance between the VDU and keyboard should be kept as short as possible to minimize evaporation from the ocular surface area by a low direction of the gaze.^[47] And 4) blink training can be beneficial.^[48]

In addition, other measures are proper lid hygiene, avoidance of eye rubbing,^[49] and proper use of personal products and medication. Eye make-up should be used with care.^[50]

Eye movement[link]

MRI scan of human eye

The visual system in the brain is too slow to process information if the images are slipping across the retina at more than a few degrees per second.^[51] Thus, for humans to be able to see while moving, the brain must compensate for the motion of the head by turning the eyes. Another complication for vision in frontal-eyed animals is the development of a small area of the retina with a very high visual acuity. This area is called the fovea centralis, and covers about 2 degrees of visual angle in people. To get a clear view of the world, the brain must turn the eyes so that the image of the object of regard falls on the fovea. Eye movements are thus very important for visual perception, and any failure to make them correctly can lead to serious visual disabilities.

Having two eyes is an added complication, because the brain must point both of them accurately enough that the object of regard falls on corresponding points of the two retinas; otherwise, double vision would occur. The movements of different body parts are controlled by striated muscles acting around joints. The movements of the eye are no exception, but they have special advantages not shared by skeletal muscles and joints, and so are considerably different.

Extraocular muscles[link]

Each eye has six muscles that control its movements: the lateral rectus, the medial rectus, the inferior rectus, the superior rectus, the inferior oblique, and the superior oblique. When the muscles exert different tensions, a torque is exerted on the globe that causes it to turn, in almost pure rotation, with only about one millimeter of translation.^[52] Thus, the eye can be considered as undergoing rotations about a single point in the center of the eye.

Rapid eye movement[link]

Rapid eye movement, or REM for short, typically refers to the sleep stage during which the most vivid dreams occur. During this stage, the eyes move rapidly. It is not in itself a unique form of eye movement.

Saccades[link]

Saccades are quick, simultaneous movements of both eyes in the same direction controlled by the frontal lobe of the brain. Some irregular drifts, movements, smaller than a saccade and larger than a microsaccade, subtend up to six minutes of arc.

Microsaccades[link]

Even when looking intently at a single spot, the eyes drift around. This ensures that individual photosensitive cells are continually stimulated in different degrees. Without changing input, these cells would otherwise stop generating output. Microsaccades move the eye no more than a total of 0.2° in adult humans.

Vestibulo-ocular reflex[link]

The vestibulo-ocular reflex is a reflex eye movement that stabilizes images on the retina during head movement by producing an eye movement in the direction opposite to head movement, thus preserving the image on the center of the visual field. For example, when the head moves to the right, the eyes move to the left, and vice versa.

Smooth pursuit movement[link]

The eyes can also follow a moving object around. This tracking is less accurate than the vestibulo-ocular reflex, as it requires the brain to process incoming visual information and supply feedback. Following an object moving at constant speed is relatively easy, though the eyes will often make saccadic jerks to keep up. The smooth pursuit movement can move the eye at up to 100°/s in adult humans.

It is more difficult to visually estimate speed in low light conditions or while moving, unless there is another point of reference for determining speed.

Optokinetic reflex[link]

The optokinetic reflex is a combination of a saccade and smooth pursuit movement. When, for example, looking out of the window at a moving train, the eyes can focus on a 'moving' train for a short moment (through smooth pursuit), until the train moves out of the field of vision. At this point, the optokinetic reflex kicks in, and moves the eye back to the point where it first saw the train (through a saccade).

Near response[link]

The adjustment to close-range vision involves three processes to focus an image on the retina.

Vergence movement[link]

The two eyes converge to point to the same object.

When a creature with binocular vision looks at an object, the eyes must rotate around a vertical axis so that the projection of the image is in the centre of the retina in both eyes. To look at an object closer by, the eyes rotate 'towards each other' (convergence), while for an object farther away they rotate 'away from each other' (divergence). Exaggerated convergence is called cross eyed viewing (focusing on the nose for example). When looking into the distance, or when 'staring into nothingness', the eyes neither converge nor diverge. Vergence movements are closely connected to accommodation of the eye. Under normal conditions, changing the focus of the eyes to look at an object at a different distance will automatically cause vergence and accommodation.

Pupil constriction[link]

Lenses cannot refract light rays at their edges as well as they can closer to the center. The image produced by any lens is therefore somewhat blurry around the edges (spherical aberration). It can be minimized by screening out peripheral light rays and looking only at the better-focused center. In the eye, the pupil serves this purpose by constricting while the eye is focused on nearby objects. In this way the pupil has a dual purpose: to adjust the eye to variations in brightness and to reduce spherical aberration.^[53]

Accommodation of the lens[link]

A change in the curvature of the lens, accommodation is carried out by the ciliary muscles surrounding the lens contracting. This narrows the diameter of the ciliary body, relaxes the fibers of the suspernsory ligament, and allows the lens to relax into a more convex shape. A more convex lens refracts light more strongly and focuses divergent light rays onto the retina allowing for closer objects to be brought into focus.^[54][55]

Effects of aging[link]

There are many diseases, disorders, and age-related changes that may affect the eyes and surrounding structures.

As the eye ages certain changes occur that can be attributed solely to the aging process. Most of these anatomic and physiologic processes follow a gradual decline. With aging, the quality of vision worsens due to reasons independent of diseases of the aging eye. While there are many changes of significance in the nondiseased eye, the most functionally important changes seem to be a reduction in pupil size and the loss of accommodation or focusing capability (presbyopia). The area of the pupil governs the amount of light that can reach the retina. The extent to which the pupil dilates decreases with age, leading to a substantial decrease in light received at the retina. In comparison to younger people, it is as though older persons are constantly wearing medium-density sunglasses. Therefore, for any detailed visually guided tasks on which performance varies with illumination, older persons require extra lighting. Certain ocular diseases can come from sexually transmitted diseases such as herpes and genital warts. If contact between eye and area of infection occurs, the STD can be transmitted to the eye.^[56]

With aging a prominent white ring develops in the periphery of the cornea- called arcus senilis. Aging causes laxity and downward shift of eyelid tissues and atrophy of the orbital fat. These changes contribute to the etiology of several eyelid disorders such as ectropion, entropion, dermatochalasis, and ptosis. The vitreous gel undergoes liquefaction (posterior vitreous detachment or PVD) and its opacities — visible as floaters — gradually increase in number.

Various eye care professionals, including ophthalmologists, optometrists, and opticians, are involved in the treatment and management of ocular and vision disorders. A Snellen chart is one type of eye chart used to measure visual acuity. At the conclusion of a complete eye examination, the eye doctor might provide the patient with an eyeglass prescription for corrective lenses. Some disorders of the eyes for which corrective lenses are prescribed include myopia (near-sightedness) which affects about one-third^{[citation needed]} of the human population, hyperopia (far-sightedness) which affects about one quarter of the population, astigmatism, and presbyopia, the loss of focusing range during aging.

Eye care professionals[link]

The human eye contains enough complexity to warrant specialized attention and care beyond the duties of a general practitioner. These specialists, or eye care professionals, serve different functions in different countries. Each eye care professional can typically be categorized into one or a multiplicity (i.e. an ophthalmologist can perform surgery; and in some instances prescribe lenses, which is a duty often performed by optometrists) of duties of the following types of professionals:

• Ophthalmologists
• Orthoptists
• Optometrists
• Opticians

See also[link]

• Eye color
• Mammalian eye
• Optometry
• Ophthalmology
• Spectral sensitivity
• Blink
• Ora serrata
• Hyaloid canal

References[link]

External links[link]

Media related to human eyes at Wikimedia Commons

• 3D Interactive Human Eye

v t e Human regional anatomy (TA A01.1) Head Forehead · Ear · Jaw · Face (Cheek, Eye, Nose, Mouth, Chin) · Occiput · Scalp · Temple Neck Throat · Adam's apple Trunk Chest (Breast) · Abdomen (Navel) · Back · Pelvis (Sex organs) Limbs Upper limb Shoulder · Axilla · Arm Elbow · Forearm Wrist · Hand · Finger (Thumb · Index · Middle · Ring · Little) Lower limb/ (see also leg) Hip · Buttocks · Thigh Knee · Calf · Crus Ankle · Heel · Foot · Toe (Hallux · Long  • Third  • Fourth  • Fifth) · Sole General anatomy: systems and organs, regional anatomy, planes and lines, superficial axial anatomy, superficial anatomy of limbs

v t e Sensory system – visual system – globe of eye (TA A15.2.1–6, TH 3.11.08.0-5, GA 10.1005) Fibrous tunic (outer) Sclera Episcleral layer • Schlemm's canal • Trabecular meshwork Cornea Limbus • layers (Epithelium, Bowman's, Stroma, Descemet's, Endothelium) Uvea/vascular tunic (middle) Choroid Capillary lamina of choroid • Bruch's membrane • Sattler's layer Ciliary body Ciliary processes • Ciliary muscle Iris Stroma • Pupil • Iris dilator muscle • Iris sphincter muscle Retina (inner) Layers Inner limiting membrane • Nerve fiber layer • Ganglion cell layer • Inner plexiform layer • Inner nuclear layer Outer plexiform layer • Outer nuclear layer External limiting membrane • Layer of rods and cones • Retinal pigment epithelium Cells Photoreceptor cells (Cone cell, Rod cell) → (Horizontal cell) → Bipolar cell → (Amacrine cell) → Retina ganglion cell (Midget cell, Parasol cell, Bistratified cell, Giant retina ganglion cells, Photosensitive ganglion cell) → Diencephalon: P cell, M cell, K cell Muller glia Other Macula (Foveola, Fovea centralis) • Optic disc (Optic cup) Anterior segment Anterior chamber • Aqueous humour • Posterior chamber • Lens (Capsule of lens, Zonule of Zinn) Posterior segment Vitreous humour Other Ocular immune system • Tapetum lucidum • Keratocytes M: EYE anat(g/a/p)/phys/devp/prot noco/cong/tumr, epon proc, drug(S1A/1E/1F/1L)

Eye
Schematic diagram of the vertebrate eye.
Compound eye of Antarctic krill

Eyes are organs that detect light and convert it into electro-chemical impulses in neurons. The simplest photoreceptors in conscious vision connect light to movement. In higher organisms the eye is a complex optical system which collects light from the surrounding environment, regulates its intensity through a diaphragm, focuses it through an adjustable assembly of lenses to form an image, converts this image into a set of electrical signals, and transmits these signals to the brain through complex neural pathways that connect the eye via the optic nerve to the visual cortex and other areas of the brain. Eyes with resolving power have come in ten fundamentally different forms, and 96% of animal species possess a complex optical system.^[1] Image-resolving eyes are present in molluscs, chordates and arthropods.^[2]

The simplest "eyes", such as those in microorganisms, do nothing but detect whether the surroundings are light or dark, which is sufficient for the entrainment of circadian rhythms.^{[citation needed]} From more complex eyes, retinal photosensitive ganglion cells send signals along the retinohypothalamic tract to the suprachiasmatic nuclei to effect circadian adjustment.

Overview[link]

Eye of the wisent,
the European bison

Complex eyes can distinguish shapes and colours. The visual fields of many organisms, especially predators, involve large areas of binocular vision to improve depth perception. In other organisms, eyes are located so as to maximise the field of view, such as in rabbits and horses, which have monocular vision.

The first proto-eyes evolved among animals 600 million years ago about the time of the Cambrian explosion.^[3] The last common ancestor of animals possessed the biochemical toolkit necessary for vision, and more advanced eyes have evolved in 96% of animal species in six of the thirty-plus^[4] main phyla.^[1] In most vertebrates and some molluscs, the eye works by allowing light to enter and project onto a light-sensitive panel of cells, known as the retina, at the rear of the eye. The cone cells (for colour) and the rod cells (for low-light contrasts) in the retina detect and convert light into neural signals for vision. The visual signals are then transmitted to the brain via the optic nerve. Such eyes are typically roughly spherical, filled with a transparent gel-like substance called the vitreous humour, with a focusing lens and often an iris; the relaxing or tightening of the muscles around the iris change the size of the pupil, thereby regulating the amount of light that enters the eye,^[5] and reducing aberrations when there is enough light.^[6]

The eyes of most cephalopods, fish, amphibians and snakes have fixed lens shapes, and focusing vision is achieved by telescoping the lens—similar to how a camera focuses.^[7]

Compound eyes are found among the arthropods and are composed of many simple facets which, depending on the details of anatomy, may give either a single pixelated image or multiple images, per eye. Each sensor has its own lens and photosensitive cell(s). Some eyes have up to 28,000 such sensors, which are arranged hexagonally, and which can give a full 360° field of vision. Compound eyes are very sensitive to motion. Some arthropods, including many Strepsiptera, have compound eyes of only a few facets, each with a retina capable of creating an image, creating vision. With each eye viewing a different thing, a fused image from all the eyes is produced in the brain, providing very different, high-resolution images.

Possessing detailed hyperspectral colour vision, the Mantis shrimp has been reported to have the world's most complex colour vision system.^[8] Trilobites, which are now extinct, had unique compound eyes. They used clear calcite crystals to form the lenses of their eyes. In this, they differ from most other arthropods, which have soft eyes. The number of lenses in such an eye varied, however: some trilobites had only one, and some had thousands of lenses in one eye.

In contrast to compound eyes, simple eyes are those that have a single lens. For example, jumping spiders have a large pair of simple eyes with a narrow field of view, supported by an array of other, smaller eyes for peripheral vision. Some insect larvae, like caterpillars, have a different type of simple eye (stemmata) which gives a rough image. Some of the simplest eyes, called ocelli, can be found in animals like some of the snails, which cannot actually "see" in the normal sense. They do have photosensitive cells, but no lens and no other means of projecting an image onto these cells. They can distinguish between light and dark, but no more. This enables snails to keep out of direct sunlight. In organisms dwelling near deep-sea vents, compound eyes have been secondarily simplified and adapted to spot the infra-red light produced by the hot vents–in this way the bearers can spot hot springs and avoid being boiled alive.^[9]

Evolution[link]

Evolution of the eye

Photoreception is phylogenetically very old, with various theories of phylogenesis.^[10] The common origin (monophyly) of all animal eyes is now widely accepted as fact. This is based upon the shared anatomical and genetic features of all eyes; that is, all modern eyes, varied as they are, have their origins in a proto-eye believed to have evolved some 540 million years ago,^[11][12][13] and the PAX6 gene is considered a key factor in this. The majority of the advancements in early eyes are believed to have taken only a few million years to develop, since the first predator to gain true imaging would have touched off an "arms race".^[14] Prey animals and competing predators alike would be at a distinct disadvantage without such capabilities and would be less likely to survive and reproduce. Hence multiple eye types and subtypes developed in parallel.

Eyes in various animals show adaptation to their requirements. For example, birds of prey have much greater visual acuity than humans, and some can see ultraviolet light. The different forms of eye in, for example, vertebrates and molluscs are often cited as examples of parallel evolution, despite their distant common ancestry.

The very earliest "eyes", called eyespots, were simple patches of photoreceptor protein in unicellular animals. In multicellular beings, multicellular eyespots evolved, physically similar to the receptor patches for taste and smell. These eyespots could only sense ambient brightness: they could distinguish light and dark, but not the direction of the light source.^[1]

Through gradual change, as the eyespot depressed into a shallow "cup" shape, the ability to slightly discriminate directional brightness was achieved by using the angle at which the light hit certain cells to identify the source. The pit deepened over time, the opening diminished in size, and the number of photoreceptor cells increased, forming an effective pinhole camera that was capable of dimly distinguishing shapes.^[15]

The thin overgrowth of transparent cells over the eye's aperture, originally formed to prevent damage to the eyespot, allowed the segregated contents of the eye chamber to specialise into a transparent humour that optimised colour filtering, blocked harmful radiation, improved the eye's refractive index, and allowed functionality outside of water. The transparent protective cells eventually split into two layers, with circulatory fluid in between that allowed wider viewing angles and greater imaging resolution, and the thickness of the transparent layer gradually increased, in most species with the transparent crystallin protein.^[16]

The gap between tissue layers naturally formed a bioconvex shape, an optimally ideal structure for a normal refractive index. Independently, a transparent layer and a nontransparent layer split forward from the lens: the cornea and iris. Separation of the forward layer again formed a humour, the aqueous humour. This increased refractive power and again eased circulatory problems. Formation of a nontransparent ring allowed more blood vessels, more circulation, and larger eye sizes.^[16]

Types of eye[link]

There are ten different eye layouts—indeed every way of capturing an optical image commonly used by human beings, with the exceptions of zoom and Fresnel lenses. Eye types can be categorised into "simple eyes", with one concave photoreceptive surface, and "compound eyes", which comprise a number of individual lenses laid out on a convex surface.^[1] Note that "simple" does not imply a reduced level of complexity or acuity. Indeed, any eye type can be adapted for almost any behaviour or environment. The only limitations specific to eye types are that of resolution—the physics of compound eyes prevents them from achieving a resolution better than 1°. Also, superposition eyes can achieve greater sensitivity than apposition eyes, so are better suited to dark-dwelling creatures.^[1] Eyes also fall into two groups on the basis of their photoreceptor's cellular construction, with the photoreceptor cells either being cilliated (as in the vertebrates) or rhabdomeric. These two groups are not monophyletic; the cnidaria also possess cilliated cells, ^[17] and some annelids possess both.^[18]

Non-compound eyes[link]

Simple eyes are rather ubiquitous, and lens-bearing eyes have evolved at least seven times in vertebrates, cephalopods, annelids, crustaceans and cubozoa.^[19]

Pit eyes[link]

Pit eyes, also known as stemma, are eye-spots which may be set into a pit to reduce the angles of light that enters and affects the eyespot, to allow the organism to deduce the angle of incoming light.^[1] Found in about 85% of phyla, these basic forms were probably the precursors to more advanced types of "simple eye". They are small, comprising up to about 100 cells covering about 100 µm.^[1] The directionality can be improved by reducing the size of the aperture, by incorporating a reflective layer behind the receptor cells, or by filling the pit with a refractile material.^[1]

Pit vipers have developed pits that function as eyes by sensing thermal infra-red radiation, in addition to their optical wavelength eyes like those of other vertebrates.

Spherical lensed eye[link]

The resolution of pit eyes can be greatly improved by incorporating a material with a higher refractive index to form a lens, which may greatly reduce the blur radius encountered—hence increasing the resolution obtainable.^[1] The most basic form, seen in some gastropods and annelids, consists of a lens of one refractive index. A far sharper image can be obtained using materials with a high refractive index, decreasing to the edges; this decreases the focal length and thus allows a sharp image to form on the retina.^[1] This also allows a larger aperture for a given sharpness of image, allowing more light to enter the lens; and a flatter lens, reducing spherical aberration.^[1] Such an inhomogeneous lens is necessary in order for the focal length to drop from about 4 times the lens radius, to 2.5 radii.^[1]

Heterogeneous eyes have evolved at least eight times: four or more times in gastropods, once in the copepods, once in the annelids and once in the cephalopods.^[1] No aquatic organisms possess homogeneous lenses; presumably the evolutionary pressure for a heterogeneous lens is great enough for this stage to be quickly "outgrown".^[1]

This eye creates an image that is sharp enough that motion of the eye can cause significant blurring. To minimise the effect of eye motion while the animal moves, most such eyes have stabilising eye muscles.^[1]

The ocelli of insects bear a simple lens, but their focal point always lies behind the retina; consequently they can never form a sharp image. This capitulates the function of the eye. Ocelli (pit-type eyes of arthropods) blur the image across the whole retina, and are consequently excellent at responding to rapid changes in light intensity across the whole visual field; this fast response is further accelerated by the large nerve bundles which rush the information to the brain.^[20] Focusing the image would also cause the sun's image to be focused on a few receptors, with the possibility of damage under the intense light; shielding the receptors would block out some light and thus reduce their sensitivity.^[20] This fast response has led to suggestions that the ocelli of insects are used mainly in flight, because they can be used to detect sudden changes in which way is up (because light, especially UV light which is absorbed by vegetation, usually comes from above).^[20]

Multiple lenses[link]

Some marine organisms bear more than one lens; for instance the copepod Pontella has three. The outer has a parabolic surface, countering the effects of spherical aberration while allowing a sharp image to be formed. Another copepod, Copilia, has two lenses in each eye, arranged like those in a telescope.^[1] Such arrangements are rare and poorly understood, but represent an interesting alternative construction. An interesting use of multiple lenses is seen in some hunters such as eagles and jumping spiders, which have a refractive cornea (discussed next): these have a negative lens, enlarging the observed image by up to 50% over the receptor cells, thus increasing their optical resolution.^[1]

Refractive cornea[link]

In the eyes of most mammals, birds, reptiles, and most other terrestrial vertebrates (along with spiders and some insect larvae) the vitreous fluid has a higher refractive index than the air.^[1] In general, the lens is not spherical. Spherical lenses produce spherical aberration. In refractive corneas, the lens tissue is corrected with inhomogeneous lens material (see Luneburg lens), or with an aspheric shape.^[1] Flattening the lens has a disadvantage; the quality of vision is diminished away from the main line of focus. Thus, animals that have evolved with a wide field-of-view often have eyes that make use of an inhomogeneous lens.^[1]

As mentioned above, a refractive cornea is only useful out of water; in water, there is little difference in refractive index between the vitreous fluid and the surrounding water. Hence creatures that have returned to the water–penguins and seals, for example–lose their highly curved cornea and return to lens-based vision. An alternative solution, borne by some divers, is to have a very strongly focusing cornea.^[1]

Reflector eyes[link]

An alternative to a lens is to line the inside of the eye with " mirrors", and reflect the image to focus at a central point.^[1] The nature of these eyes means that if one were to peer into the pupil of an eye, one would see the same image that the organism would see, reflected back out.^[1]

Many small organisms such as rotifers, copepods and platyhelminths use such organs, but these are too small to produce usable images.^[1] Some larger organisms, such as scallops, also use reflector eyes. The scallop Pecten has up to 100 millimetre-scale reflector eyes fringing the edge of its shell. It detects moving objects as they pass successive lenses.^[1]

There is at least one vertebrate, the spookfish, whose eyes include reflective optics for focusing of light. Each of the two eyes of a spookfish collects light from both above and below; the light coming from above is focused by a lens, while that coming from below, by a curved mirror composed of many layers of small reflective plates made of guanine crystals.^[21]

[edit] Compound eyes

An image of a house fly compound eye surface by using scanning electron microscope

Anatomy of the compound eye of an insect

Arthropods such as this Calliphora vomitoria fly have compound eyes

A compound eye may consist of thousands of individual photoreceptor units or ommatidia (ommatidium, singular). The image perceived is a combination of inputs from the numerous ommatidia (individual "eye units"), which are located on a convex surface, thus pointing in slightly different directions. Compared with simple eyes, compound eyes possess a very large view angle, and can detect fast movement and, in some cases, the polarisation of light.^[22] Because the individual lenses are so small, the effects of diffraction impose a limit on the possible resolution that can be obtained (assuming that they do not function as phased arrays). This can only be countered by increasing lens size and number. To see with a resolution comparable to our simple eyes, humans would require compound eyes which would each reach the size of their heads.^{[citation needed]}

Compound eyes fall into two groups: apposition eyes, which form multiple inverted images, and superposition eyes, which form a single erect image.^[23] Compound eyes are common in arthropods, and are also present in annelids and some bivalved molluscs.^[24]

Compound eyes, in arthropods at least, grow at their margins by the addition of new ommatidia.^[25]

Apposition eyes[link]

Apposition eyes are the most common form of eye, and are presumably the ancestral form of compound eye. They are found in all arthropod groups, although they may have evolved more than once within this phylum.^[1] Some annelids and bivalves also have apposition eyes. They are also possessed by Limulus, the horseshoe crab, and there are suggestions that other chelicerates developed their simple eyes by reduction from a compound starting point.^[1] (Some caterpillars appear to have evolved compound eyes from simple eyes in the opposite fashion.)

Apposition eyes work by gathering a number of images, one from each eye, and combining them in the brain, with each eye typically contributing a single point of information.

The typical apposition eye has a lens focusing light from one direction on the rhabdom, while light from other directions is absorbed by the dark wall of the ommatidium. In the other kind of apposition eye, found in the Strepsiptera, lenses are not fused to one another, and each forms an entire image; these images are combined in the brain. This is called the schizochroal compound eye or the neural superposition eye. Because images are combined additively, this arrangement allows vision under lower light levels.^[1]

Superposition eyes[link]

The second type is named the superposition eye. The superposition eye is divided into three types; the refracting, the reflecting and the parabolic superposition eye. The refracting superposition eye has a gap between the lens and the rhabdom, and no side wall. Each lens takes light at an angle to its axis and reflects it to the same angle on the other side. The result is an image at half the radius of the eye, which is where the tips of the rhabdoms are. This kind is used mostly by nocturnal insects. In the parabolic superposition compound eye type, seen in arthropods such as mayflies, the parabolic surfaces of the inside of each facet focus light from a reflector to a sensor array. Long-bodied decapod crustaceans such as shrimp, prawns, crayfish and lobsters are alone in having reflecting superposition eyes, which also have a transparent gap but use corner mirrors instead of lenses.

Parabolic superposition[link]

This eye type functions by refracting light, then using a parabolic mirror to focus the image; it combines features of superposition and apposition eyes.^[9]

Other[link]

Good fliers like flies or honey bees, or prey-catching insects like praying mantis or dragonflies, have specialised zones of ommatidia organised into a fovea area which gives acute vision. In the acute zone the eyes are flattened and the facets larger. The flattening allows more ommatidia to receive light from a spot and therefore higher resolution.

There are some exceptions from the types mentioned above. Some insects have a so-called single lens compound eye, a transitional type which is something between a superposition type of the multi-lens compound eye and the single lens eye found in animals with simple eyes. Then there is the mysid shrimp Dioptromysis paucispinosa. The shrimp has an eye of the refracting superposition type, in the rear behind this in each eye there is a single large facet that is three times in diameter the others in the eye and behind this is an enlarged crystalline cone. This projects an upright image on a specialised retina. The resulting eye is a mixture of a simple eye within a compound eye.

Another version is the pseudofaceted eye, as seen in Scutigera. This type of eye consists of a cluster of numerous ocelli on each side of the head, organised in a way that resembles a true compound eye.

The body of Ophiocoma wendtii, a type of brittle star, is covered with ommatidia, turning its whole skin into a compound eye. The same is true of many chitons. The tube feet of sea urchins contain photoreceptor proteins, which together act as a compound eye; they lack screening pigments, but can detect the directionality of light by the shadow cast by its opaque body.^[26]

Nutrients[link]

The ciliary body is triangular in horizontal section and is coated by a double layer, the ciliary epithelium. The inner layer is transparent and covers the vitreous body, and is continuous from the neural tissue of the retina. The outer layer is highly pigmented, continuous with the retinal pigment epithelium, and constitutes the cells of the dilator muscle.

The vitreous is the transparent, colourless, gelatinous mass that fills the space between the lens of the eye and the retina lining the back of the eye.^[27] It is produced by certain retinal cells. It is of rather similar composition to the cornea, but contains very few cells (mostly phagocytes which remove unwanted cellular debris in the visual field, as well as the hyalocytes of Balazs of the surface of the vitreous, which reprocess the hyaluronic acid), no blood vessels, and 98-99% of its volume is water (as opposed to 75% in the cornea) with salts, sugars, vitrosin (a type of collagen), a network of collagen type II fibres with the mucopolysaccharide hyaluronic acid, and also a wide array of proteins in micro amounts. Amazingly, with so little solid matter, it tautly holds the eye.

Relationship to life requirements[link]

Eyes are generally adapted to the environment and life requirements of the organism which bears them. For instance, the distribution of photoreceptors tends to match the area in which the highest acuity is required, with horizon-scanning organisms, such as those that live on the African plains, having a horizontal line of high-density ganglia, while tree-dwelling creatures which require good all-round vision tend to have a symmetrical distribution of ganglia, with acuity decreasing outwards from the centre.

Of course, for most eye types, it is impossible to diverge from a spherical form, so only the density of optical receptors can be altered. In organisms with compound eyes, it is the number of ommatidia rather than ganglia that reflects the region of highest data acquisition.^[1]:23-4 Optical superposition eyes are constrained to a spherical shape, but other forms of compound eyes may deform to a shape where more ommatidia are aligned to, say, the horizon, without altering the size or density of individual ommatidia.^[28] Eyes of horizon-scanning organisms have stalks so they can be easily aligned to the horizon when this is inclined, for example if the animal is on a slope.^[29] An extension of this concept is that the eyes of predators typically have a zone of very acute vision at their centre, to assist in the identification of prey.^[28] In deep water organisms, it may not be the centre of the eye that is enlarged. The hyperiid amphipods are deep water animals that feed on organisms above them. Their eyes are almost divided into two, with the upper region thought to be involved in detecting the silhouettes of potential prey—or predators—against the faint light of the sky above. Accordingly, deeper water hyperiids, where the light against which the silhouettes must be compared is dimmer, have larger "upper-eyes", and may lose the lower portion of their eyes altogether.^[28] Depth perception can be enhanced by having eyes which are enlarged in one direction; distorting the eye slightly allows the distance to the object to be estimated with a high degree of accuracy.^[9]

Acuity is higher among male organisms that mate in mid-air, as they need to be able to spot and assess potential mates against a very large backdrop.^[28] On the other hand, the eyes of organisms which operate in low light levels, such as around dawn and dusk or in deep water, tend to be larger to increase the amount of light that can be captured.^[28]

It is not only the shape of the eye that may be affected by lifestyle. Eyes can be the most visible parts of organisms, and this can act as a pressure on organisms to have more transparent eyes at the cost of function.^[28]

Eyes may be mounted on stalks to provide better all-round vision, by lifting them above an organism's carapace; this also allows them to track predators or prey without moving the head.^[9]

Visual acuity[link]

The eye of a red-tailed hawk

Visual acuity, or resolving power, is "the ability to distinguish fine detail" and is the property of cone cells.^[30] It is often measured in cycles per degree (CPD), which measures an angular resolution, or how much an eye can differentiate one object from another in terms of visual angles. Resolution in CPD can be measured by bar charts of different numbers of white/black stripe cycles. For example, if each pattern is 1.75 cm wide and is placed at 1 m distance from the eye, it will subtend an angle of 1 degree, so the number of white/black bar pairs on the pattern will be a measure of the cycles per degree of that pattern. The highest such number that the eye can resolve as stripes, or distinguish from a grey block, is then the measurement of visual acuity of the eye.

For a human eye with excellent acuity, the maximum theoretical resolution is 50 CPD^[31] (1.2 arcminute per line pair, or a 0.35 mm line pair, at 1 m). A rat can resolve only about 1 to 2 CPD.^[32] A horse has higher acuity through most of the visual field of its eyes than a human has, but does not match the high acuity of the human eye's central fovea region.^{[citation needed]}

Spherical aberration limits the resolution of a 7 mm pupil to about 3 arcminutes per line pair. At a pupil diameter of 3 mm, the spherical aberration is greatly reduced, resulting in an improved resolution of approximately 1.7 arcminutes per line pair.^[33] A resolution of 2 arcminutes per line pair, equivalent to a 1 arcminute gap in an optotype, corresponds to 20/20 (normal vision) in humans.

However, in the compound eye, the resolution is related to the size of individual ommatidia and the distance between neighbouring ommatidia. Physically these cannot be reduced in size to achieve the acuity seen with single lensed eyes as in mammals. Compound eyes have a much lower acuity than mammalian eyes.^[34]

Perception of colors[link]

"Color vision is the faculty of the organism to distinguish lights of different spectral qualities."^[35] All organisms are restricted to a small range of electromagnetic spectrum; this varies from creature to creature, but is mainly between wavelengths of 400 and 700 nm.^[36] This is a rather small section of the electromagnetic spectrum, probably reflecting the submarine evolution of the organ: water blocks out all but two small windows of the EM spectrum, and there has been no evolutionary pressure among land animals to broaden this range.^[37]

The most sensitive pigment, rhodopsin, has a peak response at 500 nm.^[38] Small changes to the genes coding for this protein can tweak the peak response by a few nm;^[2] pigments in the lens can also filter incoming light, changing the peak response.^[2] Many organisms are unable to discriminate between colours, seeing instead in shades of grey; colour vision necessitates a range of pigment cells which are primarily sensitive to smaller ranges of the spectrum. In primates, geckos, and other organisms, these take the form of cone cells, from which the more sensitive rod cells evolved.^[38] Even if organisms are physically capable of discriminating different colours, this does not necessarily mean that they can perceive the different colours; only with behavioural tests can this be deduced.^[2]

Most organisms with color vision are able to detect ultraviolet light. This high energy light can be damaging to receptor cells. With a few exceptions (snakes, placental mammals), most organisms avoid these effects by having absorbent oil droplets around their cone cells. The alternative, developed by organisms that had lost these oil droplets in the course of evolution, is to make the lens impervious to UV light — this precludes the possibility of any UV light being detected, as it does not even reach the retina.^[38]

Rods and cones[link]

The retina contains two major types of light-sensitive photoreceptor cells used for vision: the rods and the cones.

Rods cannot distinguish colours, but are responsible for low-light (scotopic) monochrome (black-and-white) vision; they work well in dim light as they contain a pigment, rhodopsin (visual purple), which is sensitive at low light intensity, but saturates at higher (photopic) intensities. Rods are distributed throughout the retina but there are none at the fovea and none at the blind spot. Rod density is greater in the peripheral retina than in the central retina.

Cones are responsible for colour vision. They require brighter light to function than rods require. In humans, there are three types of cones, maximally sensitive to long-wavelength, medium-wavelength, and short-wavelength light (often referred to as red, green, and blue, respectively, though the sensitivity peaks are not actually at these colours). The colour seen is the combined effect of stimuli to, and responses from, these three types of cone cells. Cones are mostly concentrated in and near the fovea. Only a few are present at the sides of the retina. Objects are seen most sharply in focus when their images fall on the fovea, as when one looks at an object directly. Cone cells and rods are connected through intermediate cells in the retina to nerve fibres of the optic nerve. When rods and cones are stimulated by light, the nerves send off impulses through these fibres to the brain.^[38]

Pigmentation[link]

The pigment molecules used in the eye are various, but can be used to define the evolutionary distance between different groups, and can also be an aid in determining which are closely related – although problems of convergence do exist.^[38]

Opsins are the pigments involved in photoreception. Other pigments, such as melanin, are used to shield the photoreceptor cells from light leaking in from the sides. The opsin protein group evolved long before the last common ancestor of animals, and has continued to diversify since.^[2]

There are two types of opsin involved in vision; c-opsins, which are associated with ciliary-type photoreceptor cells, and r-opsins, associated with rhabdomeric photoreceptor cells.^[39] The eyes of vertebrates usually contain cilliary cells with c-opsins, and (bilaterian) invertebrates have rhabdomeric cells in the eye with r-opsins. However, some ganglion cells of vertebrates express r-opsins, suggesting that their ancestors used this pigment in vision, and that remnants survive in the eyes.^[39] Likewise, c-opsins have been found to be expressed in the brain of some invertebrates. They may have been expressed in ciliary cells of larval eyes, which were subsequently resorbed into the brain on metamorphosis to the adult form.^[39] C-opsins are also found in some derived bilaterian-invertebrate eyes, such as the pallial eyes of the bivalve molluscs; however, the lateral eyes (which were presumably the ancestral type for this group, if eyes evolved once there) always use r-opsins.^[39] Cnidaria, which are an outgroup to the taxa mentioned above, express c-opsins - but r-opsins are yet to be found in this group.^[39] Incidentally, the melanin produced in the raticate is produced in the same fashion as that in vertebrates, suggesting the common descent of this pigment.^[39]

See also[link]

	Book: Eye
Wikipedia books are collections of articles that can be downloaded or ordered in print.

• Arthropod eye
• Cephalopod eye
• Emission theory (vision)
• Eye colour
• Eyelid
• Eye movement (sensory)
• Human eye
• Mammalian eye
• Mollusc eye
• Nictitating membrane
• Simple eye in invertebrates
• Tapetum lucidum
• Tears

References[link]

Notes[link]

Bibliography[link]

• Ali, Mohamed Ather; Klyne, M. A. (1985). Vision in Vertebrates. New York: Plenum Press. ISBN 0-306-42065-1. 

External links[link]

Wikimedia Commons has media related to: Eyes

• Evolution of the eye
• Diagram of the eye
• Webvision. The organisation of the retina and visual system. An in-depth treatment of retinal function, open to all but geared most toward graduate students.
• Eye strips images of all but bare essentials before sending visual information to brain, UC Berkeley research shows
• A collection of resources for eye anatomy.
• Anatomy of the eye - flash animated interactive.
• Eyes - Eyes are organs that detect light, and send signals along the optic nerve to the visual and other areas of the brain.
• Detection of weak optical signals by the human visual system : Perspectives in Neuroscience and in Quantum Physics
• Are Your Eyes Right February 1944 Popular Science

v t e Sensory system – visual system – globe of eye (TA A15.2.1–6, TH 3.11.08.0-5, GA 10.1005) Fibrous tunic (outer) Sclera Episcleral layer • Schlemm's canal • Trabecular meshwork Cornea Limbus • layers (Epithelium, Bowman's, Stroma, Descemet's, Endothelium) Uvea/vascular tunic (middle) Choroid Capillary lamina of choroid • Bruch's membrane • Sattler's layer Ciliary body Ciliary processes • Ciliary muscle Iris Stroma • Pupil • Iris dilator muscle • Iris sphincter muscle Retina (inner) Layers Inner limiting membrane • Nerve fiber layer • Ganglion cell layer • Inner plexiform layer • Inner nuclear layer Outer plexiform layer • Outer nuclear layer External limiting membrane • Layer of rods and cones • Retinal pigment epithelium Cells Photoreceptor cells (Cone cell, Rod cell) → (Horizontal cell) → Bipolar cell → (Amacrine cell) → Retina ganglion cell (Midget cell, Parasol cell, Bistratified cell, Giant retina ganglion cells, Photosensitive ganglion cell) → Diencephalon: P cell, M cell, K cell Muller glia Other Macula (Foveola, Fovea centralis) • Optic disc (Optic cup) Anterior segment Anterior chamber • Aqueous humour • Posterior chamber • Lens (Capsule of lens, Zonule of Zinn) Posterior segment Vitreous humour Other Ocular immune system • Tapetum lucidum • Keratocytes M: EYE anat(g/a/p)/phys/devp/prot noco/cong/tumr, epon proc, drug(S1A/1E/1F/1L)