by Dr. Gary Farr on 14 June 2002
1
Anatomy of the visual apparatus
Structures auxiliary to the eye
The eye is protected from mechanical injury by being enclosed in a socket, or orbit, which is made up of portions of several of the bones of the skull to form a four-sided pyramid the apex of which points back into the head. Thus, the floor of the orbit is made up of parts of the maxilla, zygomatic, and palatine bones, while the roof is made up of the orbital plate of the frontal bone and, behind this, by the lesser wing of the sphenoid. The optic foramen, the opening through which the optic nerve runs back into the brain and the large ophthalmic artery enters the orbit, is at the nasal side of the apex; the superior orbital fissure is a larger hole through which pass large veins and nerves. These nerves may carry nonvisual sensory messages—e.g., pain—or they may be motor nerves controlling the muscles of the eye. There are other fissures and canals transmitting nerves and blood vessels. The eyeball and its functional muscles are surrounded by a layer of orbital fat that acts much like a cushion, permitting a smooth rotation of the eyeball about a virtually fixed point, the centre of rotation. The protrusion of the eyeballs—proptosis—in exophthalmic goitre is caused by the collection of fluid in the orbital fatty tissue.
It is vitally important that the front surface of the eyeball, the cornea, remain moist. This is achieved by the eyelids, which during waking hours sweep the secretions of the lacrimal apparatus and other glands over the surface at regular intervals and which during sleep cover the eyes and prevent evaporation. The lids have the additional function of preventing injuries from foreign bodies, through the operation of the blink reflex. The lids are essentially folds of flesh covering the front of the orbit and, when the eye is open, leaving an almond-shaped aperture. The points of the almond are called canthi; that nearest the nose is the inner canthus, and the other is the outer canthus. The lid may be divided into four layers: (1) the skin, containing glands that open onto the surface of the lid margin, and the eyelashes; (2) a muscular layer containing principally the orbicularis oculi muscle, responsible for lid closure; (3) a fibrous layer that gives the lid its mechanical stability, its principal portions being the tarsal plates, one in each lid, which border directly upon the opening between the lids, called the palpebral aperture; and (4) the innermost layer of the lid, a portion of the conjunctiva. The conjunctiva is a mucous membrane that serves to attach the eyeball to the orbit and lids but permits a considerable degree of rotation of the eyeball in the orbit.
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2
Structures auxiliary to the eye
The conjunctiva lines the lids and then bends back over the surface of the eyeball, constituting an outer covering to the forward part of this and terminating at the transparent region of the eye, the cornea. The portion that lines the lids is called the palpebral portion of the conjunctiva; the portion covering the white of the eyeball is called the bulbar conjunctiva. Between the bulbar and the palpebral conjunctiva there are two loose, redundant portions forming recesses that project back toward the equator of the globe. These recesses are called the upper and lower fornices, or conjunctival sacs; it is the looseness of the conjunctiva at these points that makes movements of lids and eyeball possible. (The ophthalmologist finds the lower conjunctival sac a useful cavity in which to place drops containing drugs. He accomplishes this by merely pulling the outer lid away from the globe. The drops are retained in the cavity long enough to act directly on the cornea and to diffuse through this into the internal structures of the eye.)
The fibrous layer
The fibrous layer, which gives the lid its mechanical stability, is made up of the thick, and relatively rigid, tarsal plates, bordering directly on the palpebral aperture, and the much thinner palpebral fascia, or sheet of connective tissue; the two together are called the septum orbitale. When the lids are closed, the whole opening of the orbit is covered by this septum. Two ligaments, the medial and lateral palpebral ligaments, attached to the orbit and to the septum orbitale, stabilize the position of the lids in relation to the globe. The medial ligament is by far the stronger.
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3
The eye is kept moist by secretions of the lacrimal glands (tear glands). These almond-shaped glands under the upper lids extend inward from the outer corner of each eye. Each gland has two portions. One portion is in a shallow depression in the part of the eye socket formed by the frontal bone. The other portion projects into the back part of the upper lid. The ducts from each gland, three to 12 in number, open into the superior conjunctival fornix, or sac. From the fornix, the tears flow down across the eye and into the puncta lacrimalia, small openings at the margin of each eyelid near its inner corner. The puncta are openings into the lacrimal ducts; these carry the tears into the lacrimal sacs, the dilated upper ends of the nasolacrimal ducts, which carry the tears into the nose.
The evaporation of the tears as they flow across the eye is largely prevented by the secretion of oily and mucous material by other glands. Thus, the meibomian, or tarsal glands, consist of a row of elongated glands extending through the tarsal plates; they secrete an oil that emerges onto the surface of the lid margin and acts as a barrier for the tear fluid, which accumulates in the grooves between the eyeball and the lid barriers.
Six muscles outside the eye govern its movements. These muscles are the four rectus muscles—the inferior, medial, lateral, and superior recti—and the superior and inferior oblique muscles. The rectus muscles arise from a fibrous ring that encircles the optic nerve at the optic foramen, the opening through which the nerve passes, and are attached to the sclera, the opaque portion of the eyeball, in front of the equator, or widest part, of the eye. The superior oblique muscle arises near the rim of the optic foramen and somewhat nearer the nose than the origin of the rectus medialis. It ends in a rounded tendon that passes through a fibrous ring, the trochlea, that is attached to the frontal bone. The trochlea acts as a pulley. The tendon is attached to the sclera back of the equator of the eye.
The inferior oblique muscle originates from the floor of the orbit, passes under the eyeball like a sling, and is attached to the sclera between the attachments of the superior and lateral rectus muscles. The rectus muscles direct the gaze upward and downward and from side to side. The inferior oblique muscle tends to direct the eye upward, and the superior oblique to depress the eye; because of the obliqueness of the pull, each causes the eye to roll, and in an opposite direction.
The oblique muscles are strictly antagonistic to each other, but they work with the vertical rectus muscles in so far as the superior rectus and inferior oblique both tend to elevate the gaze and the inferior rectus and superior oblique both tend to depress the gaze. The superior and inferior recti do not produce a pure action of elevation or depression because their plane of action is not exactly vertical; in consequence, as with the obliques, they cause some degree of rolling, but by no means so great as that caused by the obliques; the direction of rolling caused by the rectus muscle is opposite to that of its synergistic oblique; the superior rectus causes the eye to roll inward, and the inferior oblique outward.
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4
Anatomy of the visual apparatus
The eye
General description
The eyeball is not a simple sphere but can be viewed as the result of fusing a small portion of a small, strongly curved sphere with a large portion of a large, not so strongly curved sphere (see figure to left). The small piece, occupying about one-sixth of the whole, has a radius of eight millimetres (0.3 inch); it is transparent and is called the cornea; the remainder, the scleral segment, is opaque and has a radius of 12 millimetres (0.5 inch). The ring where the two areas join is called the limbus. Thus, on looking directly into the eye from in front one sees the white sclera surrounding the cornea; because the latter is transparent one sees, instead of the cornea, a ring of tissue lying within the eye, the iris. 
The iris is the structure that determines the colour of the eye. The centre of this ring is called the pupil. It appears dark because the light passing into the eye is not reflected back to any great extent. By use of an ophthalmoscope, an instrument that permits the observer to illuminate the interior of the eyeball while observing through the pupil, the appearance of the interior lining of the globe can be made out; this is called the fundus; it is characterized by the large blood vessels that suppy blood to the retina; these are especially distinct as they cross over the pallid optic disk (see figure to right), or papilla, the region where the optic nerve fibres leave the globe.
The dimensions of the eye are reasonably constant, varying among individuals by only a millimetre or two; the sagittal (vertical) diameter is about 24 millimetres (about one inch) and is usually less than the transverse diameter. At birth the sagittal diameter is about 16 to 17 millimetres (about 0.65 inch); it increases rapidly to about 22.5 to 23 millimetres (about 0.89 inch) by the age of three years; between three and 13 the globe attains its full size. The weight is about 7.5 grams (.25 ounce), and its volume 6.5 millilitres (0.4 cubic inch).
The eye is made up of three coats, which enclose the optically clear aqueous humour, lens, and vitreous body. The outermost coat consists of the cornea and the sclera; the middle coat contains the main blood supply to the eye and consists, from the back forward, of the choroid, the ciliary body, and the iris. The innermost layer is the retina, lying on the choroid and receiving most of its nourishment from the vessels within the choroid, the remainder of its nourishment being derived from the retinal vessels that lie on its surface and are visible in the ophthalmoscope. The ciliary body and iris have a very thin covering, the ciliary epithelium and posterior epithelium of the iris, which is continuous with the retina.
Within the cavities formed by this triple-layered coat there are the crystalline lens, suspended by fine transparent fibres—the suspensory ligament or zonule of Zinn—from the ciliary body; the aqueous humour, a clear fluid filling the spaces between the cornea and the lens and iris; and the vitreous body, a clear jelly filling the much larger cavity enclosed by the sclera, the ciliary body, and the lens. The anterior chamber of the eye is defined as the space between the cornea and the forward surfaces of the iris and lens, while the posterior chamber is the much smaller space between the rear surface of the iris and the ciliary body, zonule, and lens; the two chambers both contain aqueous humour and are in connection through the pupil.
Outer and middle tunics of the globe
The outermost coat is made up of the cornea and the sclera. The cornea is the transparent window of the eye. It contains five distinguishable layers; the epithelium, or outer covering; Bowman's membrane; the stroma, or supporting structure; Descemet's membrane; and the endothelium, or inner lining. Up to 90 percent of the thickness of the cornea is made up of the stroma. The epithelium, which is a continuation of the epithelium of the conjunctiva, is itself made up of about six layers of cells. The superficial layer is continuously being shed, and the layers are renewed by multiplication of the cells in the innermost, or basal, layer.
The stroma appears as a set of lamellae, or plates, running parallel with the surface and superimposed on each other like the leaves of a book; between the lamellae lie the corneal corpuscles, cells that synthesize new collagen (connective tissue protein) essential for the repair and maintenance of this layer. The lamellae are made up of microscopically visible fibres that run parallel to form sheets; in successive lamellae the fibres make a large angle with each other. The lamellae in man are about 1.5 to 2.5 microns (one micron = 0.001 millimetre) thick, so that there are about 200 lamellae in the human cornea. The fibrous basis of the stroma is collagen.
Immediately above the stroma, adjacent to the epithelium, is Bowman's membrane, about eight to 14 microns thick; in the electron microscope it is evident that it is really stroma, but with the collagen fibrils not arranged in the orderly fashion seen in the rest of the stroma.
Beneath the stroma are Descemet's membrane and the endothelium. The former is about five to 10 microns thick and is made up of a different type of collagen from that in the stroma; it is secreted by the cells of the endothelium, which is a single layer of flattened cells. There is apparently no continuous renewal of these cells as with the epithelium, so that damage to this layer is a more serious matter.
The sclera is essentially the continuation backward of the cornea, the collagen fibres of the cornea being, in effect, continuous with those of the sclera. The sclera is pierced by numerous nerves and blood vessels; the largest of these holes is that formed by the optic nerve, the posterior scleral foramen. The outer two-thirds of the sclera in this region continue backward along the nerve to blend with its covering, or dural sheath—in fact, the sclera may be regarded as a continuation of the dura mater, the outer covering of the brain. The inner third of the sclera, combined with some choroidal tissue, stretches across the opening, and the sheet thus formed is perforated to permit the passage of fasciculi (bundles of fibres) of the optic nerve. This region is called the lamina cribrosa (see figure above). The blood vessels of the sclera are largely confined to a superficial layer of tissue, and these, along with the conjunctival vessels, are responsible for the bright redness of the inflamed eye. As with the cornea, the innermost layer is a single layer of endothelial cells; above this is the lamina fusca, characterized by large numbers of pigment cells.
The most obvious difference between the opaque sclera and the transparent cornea is the irregularity in the sizes and arrangement of the collagen fibrils in the sclera by contrast with the almost uniform thickness and strictly parallel array in the cornea; in addition, the cornea has a much higher percentage of mucopolysaccharide (a carbohydrate that has among its repeating units a nitrogenous sugar, hexosamine) as embedding material for the collagen fibrils. It has been shown that the regular arrangement of the fibrils is, in fact, the essential factor leading to the transparency of the cornea.
When the cornea is damaged—e.g., by a virus infection—the collagen laid down in the repair processes is not regularly arranged, with the result that an opaque patch called a leukoma, may occur.
When an eye is removed, or a man dies, the cornea soon loses its transparency, becoming hazy; this is due to the taking in of fluid from the aqueous humour, the cornea becoming thicker as it becomes hazier. The cornea can be made to reassume its transparency by maintaining it in a warm, well-aerated chamber, at about 31° C (88° F, its normal temperature); associated with this return of transparency is a loss of fluid.
Modern studies have shown that, under normal conditions, the cornea tends to take in fluid, mainly from the aqueous humour and from the small blood vessels at the limbus, but this is counteracted by a pump that expels the fluid as fast as it enters. This pumping action depends on an adequate supply of energy, and any situation that prejudices this supply causes the cornea to swell—the pump fails, or works so slowly that it cannot keep pace with the leak. Death is one cause of the failure of the pump, but this is primarily because of the loss of temperature; place the dead eye in a warm chamber and the reserves of metabolic energy it contains in the form of sugar and glycogen are adequate to keep the cornea transparent for 24 hours or more. When it is required to store corneas for grafting, as in an eye bank, it is best to remove the cornea from the globe to prevent it from absorbing fluid from the aqueous humour. The structure responsible for the pumping action is almost certainly the endothelium, so that damage to this lining can lead to a loss of transparency with swelling.
The cornea is exquisitely sensitive to pain. This is mediated by sensory nerve fibres, called ciliary nerves, that run just underneath the endothelium; they belong to the ophthalmic branch of the fifth cranial nerve, the large sensory nerve of the head. The ciliary nerves leave the globe through holes in the sclera, not in company with the optic nerve, which is concerned exclusively with responses of the retina to light.
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5
Anatomy of the visual apparatus
The eye
Outer and middle tunics of the globe
The uvea
The middle coat of the eye is called the uvea (from the Latin for “grape”) because the eye looks like a reddish-blue grape when the outer coat has been dissected away. The posterior part of the uvea, the choroid, is essentially a layer of blood vessels and connective tissue sandwiched between the sclera and the retina. The forward portion of the uvea, the ciliary body and iris, is more complex, containing as it does the ciliary muscle and the sphincter and dilator of the pupil.
The blood supply to the human eye is twofold, consisting of the retinal and uveal circulations, both of which derive from branches of the ophthalmic artery. The two systems of blood vessels differ in that the retinal vessels, which supply nutrition to the innermost layers of the retina, derive from a branch of the ophthalmic artery, called the central artery of the retina, that enters the eye with the optic nerve, while the uveal circulation, which supplies the middle and outer layers of the retina as well as the uvea, is derived from branches of the ophthalmic artery that penetrate the globe independently of the optic nerve.
The ciliary body is the forward continuation of the choroid. It is a muscular ring, triangular in horizontal section, beginning at the region called the ora serrata and ending, in front, as the root of the iris. The surface is thrown into folds, called ciliary processes, the whole being covered by the ciliary epithelium, which is a double layer of cells; the layer next to the vitreous body (see below), called the inner layer, is transparent, while the outer layer, which is continuous with the pigment epithelium of the retina, is heavily pigmented. These two layers are to be regarded embryologically as the forward continuation of the retina, which terminates at the ora serrata. Their function is to secrete the aqueous humour.
The ciliary muscle is an unstriped, involuntary, muscle concerned with alterations in the adjustments of focus—accommodation—of the optical system; the fibres run both across the muscle ring and circularly, and the effect of their contraction is to cause the whole body to move forward and to become fatter, so that the suspensory ligament that holds the lens in place is loosened.
The most forward portion of the uvea is the iris. This is the only portion that is visible to superficial inspection, appearing as a perforated disc, the central perforation, or pupil, varying in size according to the surrounding illumination and other factors. A prominent feature is the collarette at the inner edge, representing the place of attachment of the embryonic pupillary membrane that, in embryonic life, covers the pupil. As with the ciliary body, with which it is anatomically continuous, the iris consists of several layers: namely, an anterior layer of endothelium, the stroma; and the posterior iris epithelium. The stroma contains the blood vessels and the sphincter and dilator muscles; in addition, the stroma contains pigment cells that determine the colour of the eye. Posteriorly, the stroma is covered by a double layer of epithelium, the continuation forward of the ciliary epithelium; here, however, both layers are heavily pigmented and serve to prevent light from passing through the iris tissue, confining the optical pathway to the pupil. The pink iris of the albino is the result of the absence of pigment in these layers. The cells of the anterior layer of the iris epithelium have projections that become the fibres of the dilator muscle; these projections run radially, so that when they contract they pull the iris into folds and widen the pupil; by contrast, the fibres of the sphincter pupillae muscle run in a circle around the pupil, so that when they contract the pupil becomes smaller.
Usually, a baby belonging to the white races is born with blue eyes because of the absence of pigment cells in the stroma; the light reflected back from the posterior epithelium, which is blue because of scattering and selective absorption, passes through the stroma to the eye of the observer. As time goes on, pigment is deposited, and the colour changes; if much pigment is laid down the eye becomes brown or black, if little, it remains blue or gray.
The inner tunic of the globe
The inner tunic of the rear portion of the globe, as far forward as the ciliary body, is the retina, including its epithelia or coverings. These epithelia continue forward to line the remainder of the globe.
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6
The inner tunic of the globe
The epithelia
Separating the choroid (the middle tunic of the globe) from the retina proper is a layer of pigmented cells, the pigment epithelium of the retina; this acts as a restraining barrier to the indiscriminate diffusion of material from the blood in the choroid to the retina. The retina ends at the ora serrata, where the ciliary body begins (see figure to left). The pigment epithelium continues forward as a pigmented layer of cells covering the ciliary body; farther forward still, the epithelium covers the posterior surface of the iris and provides the cells that constitute the dilator muscle of this diaphragm. Next to the pigment epithelium of the retina is the neuroepithelium, or rods and cones (see below). Their continuation forward is represented by a second layer of epithelial cells covering the ciliary body, so that by the ciliary epithelium is meant the two layers of cells that are the embryological equivalent of the retinal pigment epithelium and the receptor layer (rods and cones) of the retina. This unpigmented layer of the ciliary epithelium is continued forward over the back of the iris, where it acquires pigment and is called the posterior iris epithelium.
The retina is the part of the eye that receives the light and converts it into chemical energy. The chemical energy activates nerves that conduct the messages out of the eye into the higher regions of the brain. The retina is a complex nervous structure, being, in essence, an outgrowth of the forebrain.
Ten layers of cells in the retina can be seen microscopically. In general, there are four main layers: (1) Next to the choroid is the pigment epithelium, already mentioned. (2) Beneath the epithelium is the layer of rods and cones, the light-sensitive cells. The changes induced in the rods and cones by light are transmitted to (3) a layer of neurons (nerve cells) called the bipolar cells, which are analogous to the sensory neurons that carry messages from the touch and heat receptors of the skin and transmit them to the cells of the spinal cord or the medulla (the part of the brain that is a continuation of the spinal cord). These bipolar cells connect with (4) the innermost layer of neurons, the ganglion cells; and the transmitted messages are carried out of the eye along their projections, or axons, which constitute the optic nerve fibres. Thus, the optic nerve is really a central tract, rather than a nerve, connecting two regions of the nervous system, namely, the layer of bipolar cells, and the cells of the lateral geniculate body, the latter being a visual relay station in the diencephalon (the rear portion of the forebrain).
The arrangement of the retinal cells in an orderly manner gives rise to the outer nuclear layer, containing the nuclei of the rods and cones; the inner nuclear layer, containing the nuclei and perikarya (main cell bodies outside the nucleus) of the bipolar cells, and the ganglion cell layer, containing the corresponding structures of the ganglion cells. The plexiform layers are regions in which the neurons make their interconnections. Thus, the outer plexiform layer contains the rod and cone projections terminating as the rod spherule and cone pedicle; these make connections with the dendritic processes of the bipolar cells, so that changes produced by light in the rods and cones are transmitted by way of these connections to the bipolar cells. (The dendritic process of a nerve cell is the projection that receives nerve impulses to the cell; the axon is the projection that carries impulses from the cell.) In the inner plexiform layer are the axons of the bipolar cells and the dendritic processes of the ganglion and amacrine cells (see below). The association is such as to allow messages in the bipolar cells to be transmitted to the ganglion cells, the messages then passing out along the axons of the ganglion cells as optic nerve messages.
The photosensitive cells are, in the human and in most vertebrate retinas, of two kinds, called rods and cones, the rods being usually much thinner than the cones but both being built up on the same plan. The light-sensitive pigment is contained in the outer segment, which rests on the pigment epithelium. Through the other end, called the synaptic body, effects of light are transmitted to the bipolar and horizontal cells. When examined in the electron microscope, the outer segments of the rods and cones are seen to be composed of stacks of disks, apparently made by the infolding of the limiting membrane surrounding the outer segment; the visual pigment, located on the surfaces of these disks, is thus spread over a very wide area, and this contributes to the efficiency with which light is absorbed by the visual cell.
The arrangement of the retina makes it necessary for light to pass through the layers not sensitive to light first before it reaches the light-sensitive rods and cones. The optical disadvantages of this arrangement are largely overcome by the development of the fovea centralis, a localized region of the retina, close to the optic axis of the eye, where the inner layers of the retina are absent. The result is a depression, the foveal pit, where light has an almost unrestricted passage to the light-sensitive cells. It is essentially this region of the retina that is employed for accurate vision, the eyes being directed toward the objects of regard so that their images fall in this restricted region. If the object of interest is large, so as to subtend a large angle, then the eye must move rapidly from region to region so as to bring their images successively onto the fovea; this is typically seen during reading. In the central region of the fovea there are cones exclusively; toward its edges, rods also occur, and as successive zones are reached the proportion of rods increases while the absolute density of packing of the receptors tends to decrease. Thus, the central fovea is characterized by an exclusive population of very densely packed cones; here, also, the cones are very thin and in form very similar to rods. The region surrounding the fovea is called the parafovea; it stretches about 1,250 microns from the centre of the fovea, and it is here that the highest density of rods occurs. Surrounding the parafovea, in turn, is the perifovea, its outermost edge being 2,750 microns from the centre of the fovea; here the density of cones is still further diminished, the number being only 12 per hundred microns compared with 50 per hundred microns in the most central region of the fovea. In the whole human retina there are said to be about 7,000,000 cones and from 75,000,000 to 150,000,000 rods.
The fovea is sometimes referred to as the macula lutea (“yellow spot”); actually this term defines a rather vague area, characterized by the presence of a yellow pigment in the nervous layers, stretching over the whole central retina; i.e., the fovea, parafovea, and perifovea.
The blind spot in the retina corresponds to the optic papilla, the region on the nasal side of the retina through which the optic nerve fibres pass out of the eye.
Although the rods and cones may be said to form a mosaic, the retina is not organized in a simple mosaic fashion in the sense that each rod or cone is connected to a single bipolar cell that itself is connected to a single ganglion cell. There are only about 1,000,000 optic nerve fibres, while there are at least 150,000,000 receptors, so that there must be considerable convergence of receptors on the optic pathway. This means that there will be considerable mixing of messages. Furthermore, the retina contains additional nerve cells besides the bipolar and ganglion cells; these, the horizontal and amacrine cells, operate in the horizontal direction, allowing one area of the retina to influence the activity of another. In this way, for example, the messages from one part of the retina may be suppressed by a visual stimulus falling on another, an important element in the total of messages sent to the higher regions of the brain. Finally, it has been argued that some messages may be running the opposite way; they are called centrifugal and would allow one layer of the retina to affect another, or higher regions of the brain to control the responses of the retinal neurons. In primates the existence of these centrifugal fibres has been finally disproved, but in such lower vertebrates as the pigeon, their existence is quite certain.
The pathway of the retinal messages through the brain is described later in this article; it is sufficient to state here that most of the optic nerve fibres in primates carry their messages to the lateral geniculate body, a relay station specifically concerned with vision. Some of the fibres separate from the main stream and run to a midbrain centre called the pretectal nucleus, which is a relay centre for pupillary responses to light.
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7
The transparent media
Within the cavities enclosed by the three layers of the globe described above there are the aqueous humour in the anterior and posterior chambers; the crystalline lens behind the iris; and the vitreous body, which fills the large cavity behind the lens and iris (See figure to left).
The aqueous humour
The aqueous humour is a clear colourless fluid with a chemical composition rather similar to that of blood plasma (the blood exclusive of its cells) but lacking the high protein content of the latter. Its main function is to keep the globe reasonably firm. It is secreted continuously by the ciliary body into the posterior chamber, and flows as a gentle stream through the pupil into the anterior chamber, from which it is drained by way of a channel at the limbus; that is, the juncture of the cornea and the sclera. This channel, the canal of Schlemm, encircles the cornea and connects by small connector channels to the blood vessels buried in the sclera and forming the intrascleral plexus or network. From this plexus the blood, containing the aqueous humour, passes into more superficial vessels; it finally leaves the eye in the anterior ciliary veins. The wall of the canal that faces the aqueous humour is very delicate and allows the fluid to percolate through by virtue of the relatively high pressure of the fluid within the eye. Obstruction of this exit, for example, if the iris is pushed forward to cover the wall of the canal, causes a sharp rise in the pressure within the eye, a condition that is known as {glaucoma} glaucoma. Often the obstruction is not obvious, but is caused perhaps by a hardening of the tissue just adjacent to the wall of the canal—the trabecular meshwork, in which case the rise of pressure is more gradual and insidious. Ultimately the abnormal pressure damages the retina and causes a variable degree of blindness. The normal intraocular pressure is about 15 millimetres (0.6 inch) of mercury above atmospheric pressure, so that if the anterior chamber is punctured by a hypodermic needle the aqueous humour flows out readily. Its function in maintaining the eye reasonably hard is seen by the collapse and wrinkling of the cornea when the fluid is allowed to escape. An additional function of the fluid is to provide nutrition for the crystalline lens and also for the cornea, both of which are devoid of blood vessels; the steady renewal and drainage serve to bring into the eye various nutrient substances, including glucose and amino acids, and to remove waste products of metabolism.
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8
The transparent media
The vitreous body
The vitreous body is a jelly. It is remarkable for the small amount of solid matter required to give it this semisolid structure; the solid material is made up of a form of collagen, vitrosin, and a mucopolysaccharide, hyaluronic acid. Thus, its composition is rather similar to that of the cornea, but the proportion of water is much greater, about 98 percent or more, compared with about 75 percent for the cornea. The jelly is probably secreted by certain cells of the retina. In general, the vitreous body is devoid of cells, in contrast with the lens, which is packed tight with cells. Embedded in the surface of the vitreous body, however, there is a population of specialized cells, the hyalocytes of Balazs, which may contribute to the breakdown and renewal of the hyaluronic acid. The vitreous body serves to keep the underlying retina pressed against the choroid.
The crystalline lens
The lens is a transparent body, flatter on its anterior than on its posterior surface, and suspended within the eye by the zonular fibres of Zinn attached to its equator; its anterior surface is bathed by aqueous humour, and its posterior surface by the vitreous body. The lens is a mass of tightly packed transparent fibrous cells, the lens fibres, enclosed in an elastic collagenous capsule. The lens fibres are arranged in sheets that form successive layers; the fibres run from pole to pole of the lens, the middle of a given fibre being in the equatorial region. On meridional (horizontal) section, the fibres are cut longitudinally to give an onion-scale appearance, whereas a section at right-angles to this—an equatorial section—would cut all the fibres across, and the result would be to give a honeycomb appearance. The epithelium, covering the anterior surface of the lens under the capsule, serves as the origin of the lens fibres, both during embryonic and fetal development and during infant and adult life, the lens continuing to grow by the laying down of new fibres throughout life.
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9
The work of the auxiliary structures
The protective mechanisms
The first line of protection of the eyes is provided by the lids, which prevent access of foreign bodies and assist in the lubrication of the corneal surface. Lid closure and opening are accomplished by the orbicularis oculi and levator palpebri muscles; the orbicularis oculi operates on both lids, bringing their margins into close apposition in the act of lid closure. Opening results from relaxation of the orbicularis muscle and contraction of the levator palpebri of the upper lid; the smooth muscle of the upper lid, Müller's muscle, or the superior palpebral muscle, also assists in widening the lid aperture. The lower lid does not possess a muscle corresponding to the levator of the upper lid, and the only muscle available for causing an active lowering of the lid, required during the depression of the gaze, is the inferior palpebral muscle, which is analogous to the muscle of Müller of the upper lid (called the superior palpebral muscle). This inferior palpebral muscle is so directly fused with the sheaths of the ocular muscles that it provides cooperative action, opening of the lid on downward gaze being mediated, in effect, mainly by the inferior rectus.
Innervation
The {nervous_system_cranial_d} seventh cranial nerve—the facial nerve—supplies the motor fibres for the orbicularis muscle. The levator is innervated by the third cranial nerve—the oculomotor nerve—that also innervates some of the extraocular muscles concerned with rotation of the eyeball, including the superior rectus. The smooth muscle of the eyelids and orbit is activated by the sympathetic division of the autonomic system. The secretion of adrenaline during such states of excitement as fear would also presumably cause contraction of the smooth muscle, but it seems unlikely that this would lead to the protrusion of the eyes traditionally associated with extreme fear. It is possible that the widening of the lid aperture occurring in this excited state, and dilation of the pupil, create the illusion of eye protrusion.
Blinking is normally an involuntary act, but may be carried out voluntarily. The more vigorous “full closure” of the lids involves the orbital portion of the orbicularis muscle and may be accompanied by contraction of the facial muscles that have been described as accessory muscles of blinking: namely, the corrugator supercilii, which on contraction pulls the eyebrows toward the bridge of the nose; and the procerus or pyramidalis, which pulls the skin of the forehead into horizontal folds, acting as a protection when the eyes are exposed to bright light. The more vigorous full closure may be evoked as a reflex response.
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10
The visual process
The work of the auxiliary structures
The protective mechanisms
Blink reflexes
Reflex blinking may be caused by practically any peripheral stimulus, but the two functionally significant reflexes are (1) that resulting from stimulation of the endings of the fifth cranial nerve in the cornea, lid, or conjunctiva—the sensory blink reflex, or corneal reflex—and (2) that caused by bright light—the optical blink reflex. The corneal reflex is rapid (0.1 second reflex time) and is the last to disappear in deepening anesthesia, impulses being relayed from the nucleus of the fifth nerve to the seventh cranial nerve, which transmits the motor impulses. The reflex is said to be under the control of a medullary centre. The optical reflex is slower; in man, the nervous pathway includes the visual cortex (the outer substance of the brain; the visual centre is located in the occipital—rear—lobe). The reflex is absent in children of less than nine months.
Normal rhythm
In the waking hours the eyes blink fairly regularly at intervals of two to 10 seconds, the actual rate being a characteristic of the individual. The function of this is to spread the lacrimal secretions over the cornea. It might be thought that each blink would be reflexly determined by a corneal stimulus—drying and irritation—but extensive studies indicate that this view is wrong; the normal blinking rate is apparently determined by the activity of a “blinking centre” in the globus pallidus of the caudate nucleus, a mass of nerve cells between the base and the outer substance of the brain. This is not to deny that the blink rate is modified by external stimuli.
There is a strong association between blinking and the action of the extraocular muscles. Eye movement is generally accompanied by a blink, and it is thought that this aids the eyes in changing their fixation point.
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11
The work of the auxiliary structures
The protective mechanisms
Secretion of tears
The exposed surface of the globe (eyeball) is kept moist by the tears secreted by the lacrimal apparatus, together with the mucous and oily secretions of the other secretory organs and cells of the lids and conjunctiva; these have been described earlier. The secretion produces what has been called the precorneal film, which consists of an inner layer of mucus, a middle layer of lacrimal secretion, and an outer oily film that reduces the rate of evaporation of the underlying watery layer. The normal daily (24-hour) rate of secretion has been estimated at about 0.75 to 1.1 grams (0.03–0.04 ounce avoirdupois); secretion tends to decrease with age. Chemical analysis of the tears reveals a typical body fluid with a salt concentration similar to that of blood plasma. An interesting component is lysozyme, an enzyme that has bactericidal action by virtue of its power of dissolving away the outer coats of many bacteria.
Tears are secreted reflexly in response to a variety of stimuli—e.g., irritative stimuli to the cornea, conjunctiva, nasal mucosa; hot or peppery stimuli applied to the mouth and tongue; or bright lights. In addition, tear flow occurs in association with vomiting, coughing, and yawning. The secretion associated with emotional upset is called psychical weeping. Severing of the sensory root of the trigeminal ({nervous_system_cranial_c} fifth cranial) nerve prevents all reflex weeping, leaving psychical weeping unaffected; similarly, the application of cocaine to the surface of the eye, which paralyzes the sensory nerve endings, inhibits reflex weeping, even when the eye is exposed to potent tear gases. The afferent (sensory) pathway in the reflex is thus by way of the fifth cranial, or the trigeminal nerve. The motor innervation is by way of the autonomic (involuntary) division; the parasympathetic supply derived from the facial nerve (the seventh cranial nerve) seems to have the dominant motor influence. Thus, drugs that mimic the parasympathetic, such as acetylcholine, provoke secretion, and secretion may be blocked by such typical anticholinergic drugs as atropine. Innervation of the lacrimal gland is not always complete at birth, so that the newborn infant is generally said to cry without weeping. Because absence of reflex tearing fails to produce any serious drying of the cornea, and surgical destruction of the main lacrimal gland is often without serious consequences, it seems likely that the subsidiary secretion from the accessory lacrimal glands is adequate to keep the cornea moist. The reflex secretion that produces abundant tears may be regarded as an emergency response.
A drainage mechanism for tears is necessary only during copious secretion. The mechanism, described as the lacrimal pump, consists of alternately negative and positive pressure in the lacrimal sac caused by the contraction of the orbicularis muscle during blinking.
Movements of the eyes
Because only a small portion of the retina, the fovea, is actually employed for distinct vision, it is vitally important that the motor apparatus governing the direction of gaze be extremely precise in its operation, and rapid. Thus, the gaze must shift swiftly and accurately during the process of reading. Again, if the gaze must remain fixed on a single small object—e.g., a golf ball—the eyes must keep adjusting their gaze to compensate for the continuous small movements of the head and to maintain the image exactly on the fovea. The extraocular muscles that carry out these movements are under voluntary control; thus, the direction of regard can be changed deliberately. Most of the actual movements of the eyes are carried out without awareness, however, in response to movements of the objects in the environment, or in response to movements of the head or the rest of the body, and so on. In examining the mechanisms of the eye movements, then, one must resolve them into a number of reflex responses to changes in the environment or the individual, remembering, of course, that there is an overriding voluntary control.
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12
The work of the auxiliary structures
Movements of the eyes
The axes of the eye
It is worthwhile at this point to define certain axes of the eyes employed during different types of study. The optic axis of the eye is a line drawn through the centre of the cornea and the nodal (central) point of the eye; it actually does not intersect with the retina at the centre of the fovea as might be expected, but toward the nose from this, so that there is an angle of about five degrees between (1) the visual axis—the line joining the point fixated (the point toward which the gaze is directed) and the nodal point—and (2) the optic axis.
Actions of muscles
The general modes of action of the six extraocular muscles have been described in connection with their {what_anatomy} anatomy: rotation of the eye toward the nose is carried out by the medial rectus; outward movement is by the lateral rectus. Upward movements are carried out by the combined actions of the superior rectus and the inferior oblique muscles, and downward movements by the inferior rectus and the superior oblique. Intermediate directions of gaze are achieved by combined actions of several muscles. When the two eyes act together, as they normally do, and change their direction of gaze to the left, for example, the left eye rotates away from the nose by means of its lateral rectus, while the right eye turns toward the nose by means of its medial rectus. These muscles may be considered as a linked pair; that is, when they are activated by the central nervous system this occurs conjointly and virtually automatically. This linking of the muscles of the two eyes is an important physiological feature and has still more important pathological interest in the analysis of squint, when the two eyes fail to be directed at the same point.
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13
The visual process
The work of the auxiliary structures
Movements of the eyes
Binocular movements
The binocular movements (the movements of the two eyes) fall into two classes, the conjugate movements, when both eyes move in the same direction, as in a change in the direction of gaze, and disjunctive movements, when the eyes move in opposite directions. Thus, during convergence onto a near object both eyes move toward the nose; the movement is horizontal, but disjunctive, by contrast with the conjugate movement when both eyes move, say, to the right. The disjunctive movement of convergence can be carried out voluntarily, but the act is usually brought about reflexly in response to the changed optical situation—i.e., the nearness of the object of gaze. A seesaw movement of the eyes, whereby one eye looks upward and the other downward, is possible, but not voluntarily; to achieve this a prism is placed in front on one eye so that the object seen through it appears displaced upward or downward; the other eye sees the object where it is. The result of such an arrangement is that, unless the eye with the prism in front makes an upward or downward movement, independent of the other, the images will not fall on corresponding parts of the retinas in the two eyes. Such a noncorrespondence of the retinal images causes double vision; to avoid this, there is an adjustment in the alignment of the eyes so that a seesaw movement is actually executed. In a similar way, the eyes may be made to undergo torsion, or rolling. A conjugate torsion, in which both eyes rotate about their anteroposterior (fore-and-aft) axes in the same sense, occurs naturally; for example, when the head tips toward one shoulder the eyes tend to roll in the opposite direction, with the result that the image of the visual field on the retina tends to remain vertical in spite of the rotation of the head.
Nervous control
The nerves controlling the actions of the muscles are the third, fourth, and sixth cranial nerves, with their bodies (nuclei) in the brainstem; the third, or oculomotor nerve, controls the superior and inferior recti, the medial rectus, and inferior oblique; the fourth cranial nerve, the trochlear nerve, controls the superior oblique; and the sixth, the abducens nerve, controls the lateral rectus. The nuclei of these nerves are closely associated; especially, there are connections between the nuclei of the sixth cranial nerve, controlling the lateral rectus, and the nucleus of the third, controlling the medial rectus; it is through this close relationship that the linking of the lateral rectus of one eye and the medial rectus of the other, indicated above, is achieved. Another type of linking is concerned with reciprocal inhibition; that is, when there are two antagonistic muscles, such as the medial and the lateral rectus, contraction of one is accompanied by a simultaneous inhibition of the other. Muscles show a continuous slight activity even when at rest; this keeps them taut; this action, called tonic activity, is brought about by discharges in the motor nerve to the muscle. Hence, when the agonist muscle contracts its antagonist must be inhibited.
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14
Movements of the eyes
Reflex pathways
In examining any reflex movement one must look for the sensory input—i.e., the way in which messages in sensory nerves bring about discharges in the motor nerves to the muscles; this study involves the connections of the motor nerves or nuclei with other centres of the brain.
When a subject is looking straight ahead and a bright light appears in the periphery of his field of vision, his eyes automatically turn to fix on the light; this is called the fixation reflex. The sensory pathway in the reflex arc leads as far as the cerebral cortex because removal of the {brain_b} occipital cortex (the outer brain substance at the back of the head) abolishes reflex eye movements in response to light stimuli. If the occipital cortex is stimulated electrically, movements of the eyes may be induced, and in fact one may draw a pattern of the visual field on the occipital cortex corresponding with the directions in which the gaze is turned when given points on the cortex are stimulated. This pattern corresponds with the pattern obtained by recording the visual responses to light stimuli from different parts of the visual field.
The remainder of the pathway—i.e., from the occipital cortex to the motor neurons in the brainstem—has long been considered to involve the superior colliculi as relay stations, and they certainly have such a role in lower animals; but in human beings a pathway from the cortex to the eye-muscle nuclei independent of the superior colliculi of the midbrain is now generally assumed.
Continual movements of the eyes occur even when an effort is made to maintain steady fixation of an object. Some of these movements may be regarded as manifestations of the fixation reflex; thus, the eyes tend to drift off their target, and, because of this, the fixation reflex comes into play, bringing the eyes back on target.
Experimentally, the fixation reflex can be studied by observation of the regular to-and-fro movements of the eyes as they follow a rotating drum striped in black and white. (Such movements of the eyes directed at a moving object are called optokinetic nystagmus; nystagmus itself is the involuntary movement of the eye back and forth, up and down, or in a rotatory or a mixed fashion.) While the eyes watch the moving drum, they involuntarily make a slow movement as a result of fixing their gaze on a particular stripe. At a certain point, fixation is broken off, and the eyes spring back to fix on a new stripe. Thus, the nystagmus consists of a slow movement with angular velocity equal to that of the rotation of the drum, then a fast saccade, or jump from one point of fixation to another, in the opposite direction; the process is repeated indefinitely.
Another type of nystagmus reveals the play of another set of reflexes. These are mediated by the semicircular canals—i.e., the organs of balance or the vestibular apparatus. Such a reflex may be evoked by rotating the subject in a chair at a steady speed; the eyes move slowly in the opposite direction to that of rotation and, at the end of their excursion, jump back with a fast saccade in the direction of rotation. If rotation suddenly ceases, the eyes go into a nystagmus in the opposite direction, the postrotatory nystagmus.
During rotation, certain semicircular canals are being stimulated, and the important point is that any acceleration of the head that stimulates these canals will cause reflex movements of the eyes; thus, acceleration of the head to the right causes a movement of the eyes to the left, the function of the reflex being to enable the eyes to maintain steady fixation of an object despite movements of the head. The reflex occurs even when the eyes are shut, and, when the eyes are open, it obviously cooperates with the fixation reflex in maintaining steady fixation. In many lower animals this connection between organs of balance and eyes is very rigid; thus, one may move the tail of a fish, and its eyes will move reflexly. In man, not only do the semicircular canals function in close relation to the eye muscles but so also do the gravity organ—the utricle—and the stretch receptors in the muscles of the neck. Thus, when the head is turned upward, there is a reflex tendency for the eyes to move downward, even if the eyes are shut. The actual movement is probably initiated by the reflex from the semicircular canals, which respond to acceleration, but the maintenance of the position is brought about by a reflex through the stretch of the neck muscles and also through the pull of gravity on the utricle, or otolith organ, in the inner ear.
Voluntary centre
The eyes are under voluntary control, and it is thought that the cortical area subserving voluntary eye movements is in the frontal cortex. Stimulation of this in primates causes movements of the eyes that are well coordinated, and a movement induced by this region prevails over one induced by stimulation of the occipital cortex. The existence of a separate centre in man is revealed by certain neurological disorders in which the subject is unable to fixate voluntarily but can do so reflexly; i.e., he can follow a moving light.
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15
The nature of eye movements
So far, the relation of the movements of the eyes to the requirements of the visual apparatus and their control have been touched upon. To examine the character of the movements in some detail requires rapid, accurate measurement of the movements that the eyes undergo. Modern studies of this subject employ a contact lens fitting on to the globe; on the lens is a small plane mirror, and a parallel bundle of rays is reflected off this mirror onto a moving film.
By the use of refined methods of measuring the position of the eyes at any moment, it becomes immediately evident that the eyes are never stationary for more than a fraction of a second; the movements are of three types: (1) irregular movements of high frequency (30–70 per second) and small excursions of about 20 seconds of arc; (2) flicks, or saccades, of several minutes of arc occurring at regular intervals of about one second; and between these saccades there occur (3) slow irregular drifts extending up to six minutes of arc. The saccades are corrective, serving to bring the fixation axis on the point of regard after this has drifted away from it too far, and thus are a manifestation of the fixation reflex.
The significance of these small movements during fixation was revealed by studies on the stabilized retinal image: by a suitable optical device the image of an object could be held stationary on the retina in spite of the movements of the eye. It was found that under these conditions the image would disappear within a few seconds. Thus, the movements of the eye are apparently necessary to allow the contours of the image to fall on a new set of rods and cones at repeated intervals; if this does not occur, the retina adapts to their stimulus and ceases to send messages to the central nervous system. The small flicks mentioned above are essentially the same as the larger movement made when the two eyes fixate (fix on) a light when it suddenly appears in the peripheral field; this is given the general name of the saccade, to distinguish it from the slower movements occurring during convergence and smooth following. The dynamics of the saccade have been studied in some detail by several workers. There is a reaction time of about 120 to 180 milliseconds, after which both eyes move simultaneously; there is a definite overshoot and, with an excursion of 20°, the operation is completed in about 90 milliseconds. The maximum velocity increases with the extent of the movement, being 300° per second for 10° and 500° per second for 30°. A remarkable feature is the apparent absence of significant inertia in the eyeball, so that movement is halted, not by any checking action of antagonistic muscles but simply by cessation of contraction of the agonists; thus, the movement is not ballistic. Once under way, the saccade is determined in amount, so that the subject cannot voluntarily alter its direction and extent. The control mechanism for the saccadic type of movement can be described as a sampled data system, i.e., the brain makes discontinuous samples of the position of the eyes in relation to the target and corrects the error, in contrast to a continuous feedback system that takes account of the error all the time.
The movements of the eyes when they converge onto a near object are in remarkable contrast to the saccade; the angular velocity is only about 25° per second, compared with values as high as 500° per second in the saccade. The great difference in speed suggested to two investigators that the two movements are executed by different muscle fibres. In fact, the extraocular muscles do contain two types of muscle fibre with characteristically different nerve supplies, and some recent studies tend to support this view of a dual mechanism.
If a moving light suddenly appears in the field of view, and if its rate of movement is less than about 30° per second, the response of the eyes is remarkably efficient; a saccade brings the eyes on target, and they follow the motion at almost exactly the same angular velocity as that of the target; inaccuracies in following lead to corrective saccades. When the rate of movement of the target is greater than about 30° per second, these corrective saccades become more obvious because now smooth following is not possible; the eyes make constant-velocity movements, but the velocity rarely matches that of the moving target, so that there must be frequent corrective saccades. Studies have shown that the following movements are highly integrated and must involve a continuous feedback system whereby errors are used to modify the performance. Thus, the systems for control of saccades and tracking movements are fundamentally different.
Vision suppression during a saccade
If one looks into a mirror and fixates one of one's eyes and then fixates the other, one does not see the eyes moving; and it has been argued that, during an eye movement, vision is suppressed; if vision were not suppressed, moreover, it seems likely that the images of the external world would appear smeared during a movement. Experimental studies have shown that there is, indeed, a suppression of vision during a saccade.
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16
The work of the optical lens system
Refraction by cornea and lens
The optical system of the eye is such as to produce a reduced inverted image of the visual field on the retina; the system behaves as a convex lens but is, in fact, much more complex, refraction taking place not at two surfaces, as in a lens, but at four separate surfaces—at the anterior and the posterior surfaces of the cornea and of the crystalline lens. Each of these surfaces is approximately spherical, and at each optical interface—e.g., between air and the anterior surface of the cornea—the bending of a ray of light is toward the axis, so that, in effect, there are four surfaces tending to make rays of light converge on each other. If the rays of light falling on the cornea are parallel—i.e., if they come from a distant point—the net effect of this series of refractions at the four surfaces is to bring these rays to a point focus of the optical system, which in the normal, or emmetropic, eye corresponds with the retina. The greatest change of direction, or bending of the rays, occurs where the difference of refractive index is greatest, and this is when light passes from air into the cornea, the refractive index of the corneal substance being 1.3376; the refractive indices of the cornea and aqueous humour are not greatly different, that of the aqueous humour being 1.336 (as is that of the vitreous); thus, the bending, as the rays meet the concave posterior surface of the cornea and emerge into a medium of slightly less refractive index, is small. The lens has a greater refractive index than that of its surrounding aqueous humour and vitreous body, 1.386 to 1.406, so that its two surfaces contribute to convergence, the posterior surface normally more than the anterior surface because of its greater curvature (smaller radius).
Normal sightedness and near- and farsightedness
In contrast to the focussing of the normal (emmetropic) eye, in which the image of the visual field is focussed on the retina, the image may be focussed in front of the retina (nearsightedness, or myopia), or behind the retina (farsightedness or hyperopia). In myopia the vision of distant objects is not distinct because the image of a distant point falls within the vitreous and the rays spread out to form a blur circle on the retina instead of a point. In this condition the eye is said to have too great dioptric (refractive) power for its length. When the focus falls behind the retina, the image of the distant point is again a circle on the retina; and the farsighted eye is said to have too little dioptric power. The important point to appreciate is that emmetropia, or normal sight, requires that the focal power of the dioptric system be matched to the axial length of the eye; it certainly is remarkable that emmetropia is indeed the most common condition when it is appreciated that just one millimetre of error in the matching of axial length with focal length would cause a person to require a spectacle correction. In general, however, the effects of variations in dimensions tend to compensate each other. Thus, for example, an unusually large eye might, at first thought, be expected to be myopic, but a large eye tends to be associated with a large radius of curvature of the cornea, and this would reduce the power—i.e., increase the focal length—and so an unusually large eye is not necessarily a myopic one.
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17
Effects of accommodation
The image of an object brought close to the eye would be formed behind the retina if there were no change in the focal length of the eye. This change to bring the image of an object upon the retina is called accommodation. The point nearer than which accommodation is no longer effective is called the near point of accommodation. In very young people, the near point of accommodation is quite close to the eye, namely about seven centimetres (about three inches) in front at 10 years old; at 40 years the distance has increased to about 16 centimetres (about 6 inches), and at 60 years it is 100 centimetres or one metre (39 inches). Thus, a 60-year-old would not be able to read a book held at the convenient distance of about 40 centimetres (16 inches), and the extra power required would have to be provided by convex lenses in front of the eye, an arrangement called the presbyopic correction.
Mechanism of accommodation
It is essentially an increase in curvature of the anterior surface of the lens that is responsible for the increase in power involved in the process of accommodation. A clue to the way in which this change in shape takes place is given by the observation that a lens that has been taken out of the eye is much rounder and fatter than one within the eye; thus, its attachments by the zonular fibres to the ciliary muscle within the eye preserve the unaccommodated or flattened state of the lens; and modern investigations leave little doubt that it is the pull of the zonular fibres on the elastic capsule of the lens that holds the anterior surface relatively flat. When these zonular fibres are loosened, the elastic tension in the capsule comes into play and remolds the lens, making it smaller and thicker. Thus, the physiological problem is to find what loosens the zonular fibres during accommodation. The ciliary muscle has been described earlier, and it has been shown that the effect of contracting its fibres is, in general, to pull the whole ciliary body forward and to move the anterior region toward the axis of the eye by virtue of the sphincter action of the circular fibres. Both of these actions will slacken the zonular fibres and therefore allow the change in shape. As to why it is the anterior surface that changes most is not absolutely clear, but it is probably a characteristic of the capsule rather than of the underlying lens tissue. Defective accommodation in presbyopia is not due to a failure of the ciliary muscle but rather to a hardening of the substance of the lens with age to the point that readjustments of its shape become ever more difficult.
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18
Nerve action
Accommodation is an involuntary reflex act, and the ciliary muscle belongs to the smooth involuntary class. Appropriate to this, the innervation is through the autonomic system, the parasympathetic nerve cells belonging to the oculomotor nerve (the third cranial nerve) occupying a special region of the nucleus in the midbrain called the Edinger-Westphal nucleus; the fibres have a relay point in the ciliary ganglion in the eye socket, and the postganglionic fibres enter the eye as the short ciliary nerves. The stimulus for accommodation is the nearness of the object, but the manner in which this nearness is translated into a stimulus is not clear. Thus, the fact that the image is blurred is not sufficient to induce accommodation; the eye has some power of discriminating whether the blurredness is due to an object being too far away or too close, so that something more than mere blurredness is required.
The pupil
The amount of light entering the eye is restricted by the aperture in the iris, the pupil.
When a person is in a dark room his pupil is large, perhaps eight millimetres (0.3 inch) in diameter, or more. When the room is lighted there is an immediate constriction of the pupil, the light reflex; this is bilateral, so that even if only one eye is exposed to the light both pupils contract to nearly the same extent. After a time the pupils expand even though the bright light is maintained, but the expansion is not large. The final state is determined by the actual degree of illumination; if this is high, then the final state may be a diameter of only about three to four millimetres (about 0.15 inch); if it is not so high, then the initial constriction may be nearly the same, but the final state may be with a pupil of four to five millimetres (about 0.18 inch). During this steady condition, the pupils do not remain at exactly constant size; there is a characteristic oscillation in size that, if exaggerated, is called hippus.
A pupillary constriction will also occur when a person looks at a near object—the near reflex. Thus, accommodation and pupillary constriction occur together reflexly and are excited by the same stimulus. The function of the pupil is clearly that of controlling the amount of light entering the eye, and hence the light reflex. The constriction occurring during near vision suggests other functions, too; thus, the aberrations of the eye (failure of some refracted rays to focus on the retina) are decreased by reducing the aperture of its optical system. In the dark, aberrations are of negligible significance, so that a person is concerned only with allowing as much light into the eye as possible; in bright light high visual acuity is usually required, and this means reducing the aberrations. The depth of focus of the optical system is increased when the aperture is reduced, and the near reflex is probably concerned with increasing depth of focus under these conditions.
Dilation of the pupil occurs as a result of strong psychical stimuli and also when any sensory nerve is stimulated; dilation thus occurs in extreme fear and in pain.
The muscles of the iris have been described earlier. It is clear from their general features that constriction of the pupil is brought about by shortening of the circular ring of fibres—the sphincter; dilation is brought about by shortening of the radially oriented fibres. The sphincter is innervated by parasympathetic fibres of the oculomotor nerve, with their cell bodies in the Edinger-Westphal nucleus, as are the nerve cells controlling accommodation; thus, the close association between the accommodation and pupillary reflexes is reflected in a close anatomical contiguity of their motor nerve cells.
The sensory pathway in the light reflex involves the rods and cones, bipolar cells, and ganglion cells. As indicated earlier, a relay centre for pupillary responses to light is the pretectal nucleus in the midbrain. There is a partial crossing-over of the fibres of the pretectal nerve cells so that some may run to the motor nerve cells in the Edinger-Westphal nucleus of both sides of the brain, and it is by this means that illumination of one eye affects the other. The Edinger-Westphal motor neurons have a relay point in the ciliary ganglion, a group of nerve cells in the eye socket, so that its electrical stimulation causes both accommodation and pupillary constriction; similarly, application of a drug, such as pilocarpine, to the cornea will cause a constriction of the pupil and also a spasm of accommodation; atropine, by paralyzing the nerve supply, causes dilation of the pupil and paralysis of accommodation (cycloplegia).
The dilator muscle of the iris is activated by sympathetic nerve fibres. Stimulation of the sympathetic nerve in the neck causes a powerful dilation of the iris; again, the influx of adrenalin into the blood from the adrenal glands during extreme excitement results in pupillary dilation.
Many involuntary muscles receive a double innervation, being activated by one type of nerve supply and inhibited by the other; modern experimentation indicates that the iris muscles are no exception, so that the sphincter has an inhibitory sympathetic nerve supply, while the dilator has a parasympathetic (cholinergic) inhibitor. Thus, a drug like pilocarpine not only activates the constrictor muscle but actively inhibits the dilator. A similar double innervation has been described for the ciliary muscle. In general, any change in pupillary size results from a reciprocal innervation of dilator and constrictor; thus, activation of the constrictor is associated with inhibition of the dilator and vice versa.
The near response
In general, as has been indicated, pupillary constriction and accommodation occur together, in response to the same stimulus; a third element in this near response is, of course, the convergence (turning in) of the eyes, mediated by voluntary muscles, the medial recti. Experimentally, it is often possible to separate these activities, in the sense that one may cause convergence without accommodation by placing appropriate prisms in front of the eyes; or one may cause accommodation without convergence by placing diverging lenses in front of the eyes. There are many experiments that show that accommodation and convergence are neurologically linked to some extent, however.
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