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Editor-In-Chief: C. Michael Gibson, M.S., M.D. [1]


A human eye

Eyes are organs that detect light. Different kinds of light-sensitive organs are found in a variety of animals. The simplest eyes do nothing but detect whether the surroundings are light or dark, which is sufficient for the entrainment of circadian rhythms but hardly can be called vision. More complex eyes can distinguish shapes and colors. The visual fields of some such complex eyes largely overlap, to allow better depth perception (binocular vision), as in humans; and others are placed so as to minimize the overlap, such as in rabbits and chameleons.

In the human eye, light enters the pupil and is focused on the retina by the lens. Light-sensitive nerve cells called rods (for brightness) and cones (for color) react to the light. They interact with each other and send messages to the brain that indicate brightness, color, and contour.

The first proto-eyes evolved among animals 540 million years ago. Almost all animals have eyes, or descend from animals that did.

In most vertebrates and some mollusks, the eye works by allowing light to enter it 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. 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 which regulates the intensity of the light that enters the eye. The eyes of cephalopods, fish, amphibians and snakes usually have fixed lens shapes, and focusing vision is achieved by telescoping the lens—similar to how a camera focuses.

Compound eyes are found among the arthropods and are composed of many simple facets which give a pixelated image (not multiple images, as is often believed). 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-degree 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 multiple-image vision. With each eye viewing a different angle, a fused image from all the eyes is produced in the brain, providing very wide-angle, high-resolution images.

Possessing detailed hyperspectral color vision, the Mantis shrimp has been reported to have the world's most complex color vision system.[1] 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.

Some of the simplest eyes, called ocelli, can be found in animals like snails, who 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. Jumping spiders have simple eyes that are so large, supported by an array of other, smaller eyes, that they can get enough visual input to hunt and pounce on their prey. Some insect larvae, like caterpillars, have a different type of simple eye (stemmata) which gives a rough image.

Evolution of eyes

File:Diagram of eye evolution.svg

Biologists use the theory of evolution to explain the origin and development of eyes, as well as of organs in general.

The common origin (monophyly) of all animal eyes is established by shared anatomical and genetic features of all eyes; that is, all modern eyes, varied as they are, have their origins in a proto-eye evolved some 540 million years ago.[2][3][4] The majority of the advancements in early eyes are believed to have taken only a few million years to develop, as the first predator to gain true imaging would have touched off an "arms race",[5] or rather, a phylogenetic radiation from the species with that first proto-eye, among the descendents of which, there may well have been an "arms race". Prey animals and competing predators alike would be forced to rapidly match or exceed any such capabilities to survive. Hence multiple eye types and subtypes developed in parallel.

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

The earliest eyes, called "eyespots", were simple patches of photoreceptor cells, or light-sensitive proteins in unicellular organisms, 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 lightsource.[6] This gradually changed as the eyespot depressed into a shallow "cup" shape, granting the ability to slightly discriminate directional brightness 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 slightly distinguishing dim shapes.[7]

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 specialize into a transparent humour that optimized color 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.[8]

Anatomy of the mammalian eye

Three layers

The structure of the mammalian eye can be divided into three main layers or tunics whose names reflect their basic functions: the fibrous tunic, the vascular tunic, and the nervous tunic.[9][10][11]

  • The fibrous tunic, also known as the tunica fibrosa oculi, is the outer layer of the eyeball consisting of the cornea and sclera.[12] The sclera gives the eye most of its white color. It consists of dense connective tissue filled with the protein collagen to both protect the inner components of the eye and maintain its shape.[13]
  • The vascular tunic, also known as the tunica vasculosa oculi, is the middle vascularized layer which includes the iris, ciliary body, and choroid.[12][14][15] The choroid contains blood vessels that supply the retinal cells with necessary oxygen and remove the waste products of respiration. The choroid gives the inner eye a dark color, which prevents disruptive reflections within the eye.
  • The nervous tunic, also known as the tunica nervosa oculi, is the inner sensory which includes the retina.[12][15] The retina contains the photosensitive rod and cone cells and associated neurons. To maximise vision and light absorption, the retina is a relatively smooth (but curved) layer. It has two points at which it is different; the fovea and optic disc. The fovea is a dip in the retina directly opposite the lens, which is densely packed with cone cells. It is largely responsible for color vision in humans, and enables high acuity, such as is necessary in reading. The optic disc, sometimes referred to as the anatomical blind spot, is a point on the retina where the optic nerve pierces the retina to connect to the nerve cells on its inside. No photosensitive cells exist at this point, it is thus "blind". In addition to the rods and cones, a small proportion (about 2% in humans) of the ganglion cells in the retina are photosensitive through the pigment melanopsin. They are generally most excitable by blue light, about 470 nm. Their information is sent to the SCN (suprachiasmatic nuclei), not to the visual center, through the retinohypothalamic tract, not via the optic nerve. It is these light signals which regulate circadian rhythms in mammals and several other animals. Many, but not all, totally blind individuals have their circadian rhythms adjusted daily in this way.

Anterior and posterior segments

The mammalian eye can also be divided into two main segments: the anterior segment and the posterior segment.[16]

Posterior segment

Diagram of a human eye; note that not all eyes have the same anatomy as a human eye.

The posterior segment is the back two-thirds of the eye that includes the anterior hyaloid membrane and all structures behind it: the vitreous humor, retina, choroid, and optic nerve.[17] On the other side of the lens is the second humour, the vitreous humour, which is bounded on all sides: by the lens, ciliary body, suspensory ligaments and by the retina. It lets light through without refraction, helps maintain the shape of the eye and suspends the delicate lens. In some animals, the retina contains a reflective layer (the tapetum lucidum) which increases the amount of light each photosensitive cell perceives, allowing the animal to see better under low light conditions.

File:Focus in an eye.svg

Extraocular anatomy

In many species, the eyes are inset in the portion of the skull known as the orbits or eyesockets. This placement of the eyes helps to protect them from injury.

In humans, the eyebrows redirect flowing substances (such as rainwater or sweat) away from the eye. Water in the eye can alter the refractive properties of the eye and blur vision. It can also wash away the tear fluid—along with it the protective lipid layer—and can alter corneal physiology, due to osmotic differences between tear fluid and freshwater. Osmotic effects are made apparent when swimming in freshwater pools, as the osmotic gradient draws "pool water" into the corneal tissue (the pool water is hypotonic), causing edema, and subsequently leaving the swimmer with "cloudy" or "misty" vision for a short period thereafter. The edema can be reversed by irrigating the eye with hypertonic saline which osmotically draws the excess water out of the eye.

In many animals, including humans, eyelids wipe the eye and prevent dehydration. They spread tears on the eyes, which contains substances which help fight bacterial infection as part of the immune system. Some aquatic animals have a second eyelid in each eye which refracts the light and helps them see clearly both above and below water. Most creatures will automatically react to a threat to its eyes (such as an object moving straight at the eye, or a bright light) by covering the eyes, and/or by turning the eyes away from the threat. Blinking the eyes is, of course, also a reflex.

In many animals, including humans, eyelashes prevent fine particles from entering the eye. Fine particles can be bacteria, but also simple dust which can cause irritation of the eye, and lead to tears and subsequent blurred vision.

Cytology

The structure of the mammalian eye owes itself completely to the task of focusing light onto the retina. This light causes chemical changes in the photosensitive cells of the retina, the products of which trigger nerve impulses which travel to the brain.

The retina contains two forms of photosensitive cells important to vision—rods and cones. Though structurally and metabolically similar, their function is quite different. Rod cells are highly sensitive to light, allowing them to respond in dim light and dark conditions; however, they cannot detect color differences. These are the cells that allow humans and other animals to see by moonlight, or with very little available light (as in a dark room). Cone cells, conversely, need high light intensities to respond and have high visual acuity. Different cone cells respond to different wavelengths of light, which allows an organism to see color. The shift from cone vision to rod vision is why the darker conditions become, the less color objects seem to have.

The differences between rods and cones are useful; apart from enabling sight in both dim and light conditions, they have further advantages. The fovea, directly behind the lens, consists of mostly densely-packed cone cells. The fovea gives humans a highly detailed central vision, allowing reading, bird watching, or any other task which primarily requires staring at things. Its requirement for high intensity light does cause problems for astronomers, as they cannot see dim stars, or other celestial objects, using central vision because the light from these is not enough to stimulate cone cells. Because cone cells are all that exist directly in the fovea, astronomers have to look at stars through the "corner of their eyes" (averted vision) where rods also exist, and where the light is sufficient to stimulate cells, allowing an individual to observe faint objects.

Rods and cones are both photosensitive, but respond differently to different frequencies of light. They contain different pigmented photoreceptor proteins. Rod cells contain the protein rhodopsin and cone cells contain different proteins for each color-range. The process through which these proteins go is quite similar — upon being subjected to electromagnetic radiation of a particular wavelength and intensity, the protein breaks down into two constituent products. Rhodopsin, of rods, breaks down into opsin and retinal; iodopsin of cones breaks down into photopsin and retinal. The opsin in each opens ion channels on the cell membrane, which leads to hyperpolarization; this hyperpolarization of the cell leads to a release of transmitter molecules at the synapse.

Differences between the rhodopsin and the iodopsins is the reason why cones and rods enable organisms to see in dark and light conditions — each of the photoreceptor proteins requires a different light intensity to break down into the constituent products. Further, synaptic convergence means that several rod cells are connected to a single bipolar cell, which then connects to a single ganglion cell by which information is relayed to the visual cortex. This convergence is in direct contrast to the situation with cones, where each cone cell is connected to a single bipolar cell. This divergence results in the high visual acuity, or the high ability to distinguish detail, of cone cells compared to rods. If a ray of light were to reach just one rod cell, the cell's reponse may not be enough to hyperpolarize the connected bipolar cell. But because several "converge" onto a bipolar cell, enough transmitter molecules reach the synapses of the bipolar cell to hyperpolarize it.

Furthermore, color is distinguishable due to the different iodopsins of cone cells; there are three different kinds, in normal human vision, which is why we need three different primary colors to make a color space.

Acuity

Visual acuity 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 gray block, is then the measurement of visual acuity of the eye.

For a human eye with excellent acuity, the maximum theoretical resolution would be 50 CPD[18] (1.2 minute of arc per line pair, or a 0.35 mm line pair, at 1 m). However, the eye can only resolve a contrast of 5%. Taking this into account, the eye can resolve a maximum resolution of 37 CPD, or 1.6 minute of arc per line pair (0.47 mm line pair, at 1 m). [19] A rat can resolve only about 1 to 2 CPD.[20] 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.

Equivalent resolution

A maximum resolution of the human eye in good light of 1.6 minute of arc per line pair will correspond to 1.25 lines per minute of arc. Assuming two pixels per line pair (one pixel per line) and a square field of 120 degrees, this would be equivalent to approximately 120×60×1.25 = 9000 pixels in each of the X and Y dimensions, or about 81 megapixels.

However, the human eye itself has only a small spot of sharp vision in the middle of the retina, the fovea centralis, the rest of the field of view being progressively lower resolution as it gets further from the fovea. The angle of the sharp vision being just a few degrees in the middle of the view, the sharp area thus barely achieves even a single megapixel resolution. The experience of wide sharp human vision is in fact based on turning the eyes towards the current point of interest in the field of view, the brain thus perceiving an observation of a wide sharp field of view.

The narrow beam of sharp vision is easy to test by putting a fingertip on a newspaper and trying to read the text while staring at the fingertip — it is very difficult to read text that's just a few centimeters away from the fingertip.

Spectral response

Human eyes respond to light with wavelength in the range of approximately 400 to 700 nm. Other animals have other ranges, with many such as birds including a significant ultraviolet (shorter than 400 nm) response.

Dynamic range

The retina has a static contrast ratio of around 100:1 (about 6 1/2 stops). As soon as the eye moves (saccades) it re-adjusts its exposure both chemically and by adjusting the iris. 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 stops) is possible. The process is nonlinear and multifaceted, so an interruption by light nearly 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 alcohol or tobacco.

Eye movement

MRI scan of human eye

The visual system in the brain is too slow to process that information if the images are slipping across the retina at more than a few degrees per second.[21] 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, 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

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.[22] Thus, the eye can be considered as undergoing rotations about a single point in the center of the eye. Once the human eye sustains damage to the optic nerve, the impulses will not be taken to the brain. Eye transplants can happen but the person receiving the transplant will not be able to see. As for the optic nerve, once it is damaged it cannot be fixed.

Rapid eye movement

Rapid eye movement, or REM for short, typically refers to the stage during sleep 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

Saccades are quick, simultaneous movements of both eyes in the same direction controlled by the frontal lobe of the brain.

Microsaccades

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

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

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

The optokinetic reflex is a combination of a saccade and smooth pursuit movement. When, for example, looking out of the window in a moving train, the eyes can focus on a 'moving' tree for a short moment (through smooth pursuit), until the tree 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 tree (through a saccade).

Vergence movement

File:Stereogram Tut Eye Convergence.png

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.

Accommodation

To see clearly, the lens will be pulled flatter or allowed to regain its thicker form.

Diseases, disorders, and age-related changes

The stye is a common irritating inflammation of the eyelid.

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 aging eye diseases. 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 also decreases with age. Because of the smaller pupil size, older eyes receive much less light at the retina. In comparison to younger people, it is as though older persons wear medium-density sunglasses in bright light and extremely dark glasses in dim light. 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 will be transmitted to the eye.[23]

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 an eye examination, an eye doctor may provide the patient with an eyeglass prescription for corrective lenses

Eye injury/safety

Accidents involving common household products cause 125,000 eye injuries each year in the U.S.[24] More than 40,000 people a year suffer eye injuries while playing sports.[24] Sports-related eye injuries occur most frequently in baseball, basketball and racquet sports.[24]

Occupational eye injury

Each day about 2000 U.S. workers have a job-related eye injury that requires medical treatment.[25] About one third of the injuries are treated in hospital emergency departments and more than 100 of these injuries result in one or more days of lost work.[25] The majority of these injuries result from small particles or objects striking or abrading the eye. Examples include metal slivers, wood chips, dust, and cement chips that are ejected by tools, wind blown, or fall from above a worker. Some of these objects, such as nails, staples, or slivers of wood or metal penetrate the eyeball and result in a permanent loss of vision. Large objects may also strike the eye/face causing blunt force trauma to the eyeball or eye socket. Chemical burns to one or both eyes from splashes of industrial chemicals or cleaning products are common. Thermal burns to the eye occur as well. Among welders, their assistants, and nearby workers, UV radiation burns (welder’s flash) routinely damage workers’ eyes and surrounding tissue. In addition to common eye injuries, health care workers, laboratory staff, janitorial workers, animal handlers, and other workers may be at risk of acquiring infectious diseases via ocular exposure.[25]


See also

References

  1. http://www.nwf.org/nationalwildlife/article.cfm?issueID=77&articleID=1114
  2. Halder, G., Callaerts, P. and Gehring, W.J. (1995). "New perspectives on eye evolution." Curr. Opin. Genet. Dev. 5 (pp. 602–609).
  3. Halder, G., Callaerts, P. and Gehring, W.J. (1995). "Induction of ectopic eyes by targeted expression of the eyeless gene in Drosophila". Science 267 (pp. 1788–1792).
  4. Tomarev, S.I., Callaerts, P., Kos, L., Zinovieva, R., Halder, G., Gehring, W., and Piatigorsky, J. (1997). "Squid Pax-6 and eye development." Proc. Natl. Acad. Sci. USA, 94 (pp. 2421–2426).
  5. Conway-Morris, S. (1998). The Crucible of Creation. Oxford: Oxford University Press.
  6. Land, M.F. and Fernald, Russell D. (1992). "The evolution of eyes." Annu Rev Neurosci 15 (pp. 1–29).
  7. Eye-Evolution?
  8. Fernald, Russell D. (2001). The Evolution of Eyes: Where Do Lenses Come From? Karger Gazette 64: "The Eye in Focus".
  9. "The Eye." Accessed October 23, 2006.
  10. "General Anatomy of the Eye." Accessed October 23, 2006.
  11. "Eye Anatomy and Function." Accessed October 23, 2006.
  12. 12.0 12.1 12.2 Cline D; Hofstetter HW; Griffin JR. Dictionary of Visual Science. 4th ed. Butterworth-Heinemann, Boston 1997. ISBN 0-7506-9895-0
  13. http://www.bartleby.com/107/225.html
  14. Cassin, B. and Solomon, S. Dictionary of Eye Terminology. Gainsville, Florida: Triad Publishing Company, 1990.
  15. 15.0 15.1 "Medline Encyclopedia: Eye." Accessed October 25, 2006.
  16. http://www.e-sunbear.com/anatomy_02.html
  17. Posterior segment anatomy
  18. John C. Russ (2006). The Image Processing Handbook. CRC Press. ISBN 0849372542. 
  19. Optical System Design. McGraw-Hill Professional. 2000. ISBN 0071349162.  Text " Steve Chapman (editor) " ignored (help)
  20. Curtis D. Klaassen (2001). Casarett and Doull's Toxicology: The Basic Science of Poisons. McGraw-Hill Professional. ISBN 0071347216. 
  21. Westheimer, Gerald & McKee, Suzanne P.; "Visual acuity in the presence of retinal-image motion". Journal of the Optical Society of America 1975 65(7), 847-50.
  22. Roger H.S. Carpenter (1988); Movements of the Eyes (2nd ed.). Pion Ltd, London. ISBN 0-85086-109-8.
  23. AgingEye Times
  24. 24.0 24.1 24.2 "Eye Safety Prevent Eye Injuries at Home, at Work and at Play!". Prevent Blindness America. Retrieved 2007-10-23. 
  25. 25.0 25.1 25.2 "NIOSH Eye Safety". United States National Institute for Occupational Safety and Health. Retrieved 2007-10-23. 
  • "Anatomy". History of Ophthalmology.  Unknown parameter |accessdaymonth= ignored (help); Unknown parameter |accessyear= ignored (|access-date= suggested) (help)
  • Kandel ER, Schwartz JH, Jessell TM. Principles of Neural Science, 4th ed. McGraw-Hill, New York (2000). ISBN 0-8385-7701-6

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