Pengantar Fisologi Penglihatan

M. Djauhari Widjajakusumah Departemen Fisiologi Fakultas Kedokteran Unversitas Indonesia

Dee Silverthorn

Penglihatan

INTRODUCTION
o

The eyes are complex sense organs that have evolved from primitive light-sensitive spots on the surface of invertebrates. Within its protective casing, each eye has o a layer of receptors o a lens system that focuses light on these receptors o a system of nerves that conducts impulses from the receptors to the brain.

The way these components operate to set up conscious visual images is the subject of this chapter.
Ganongs Review of Med Physiol, 23rd ed

ANATOMIC CONSIDERATIONS
Ganongs Review of Med Physiol 22nd ed Fig 8-1

Figure 81.

ANATOMIC CONSIDERATIONS

ANATOMIC CONSIDERATIONS
o The sclera is modified anteriorly to form the transparent cornea, through which light rays enter the eye. o The choroid, is a layer inside the sclera that contains many of the blood vessels that nourish the structures in the eyeball. o The retina is lining the posterior two thirds of the choroid, the neural tissue containing the receptor cells. o The crystalline lens is a transparent structure held in place by a circular lens suspensary ligament (zonule). The zonule is attached to the thickened anterior part of the choroid, the ciliary body that contains circular muscle fibers and longitudinal muscle fibers that attach near the corneoscleral junction. o In front of the lens is the pigmented and opaque iris, the colored portion of the eye. Contains circular muscle fibers that constrict and radial fibers that dilate the pupil.
Ganongs Review of Med Physiol, 23rd ed

ANATOMIC CONSIDERATIONS
o

The space between the lens and the retina is filled primarily with a clear gelatinous material called the vitreous (vitreous humor)

Aqueous humor, a clear liquid that nourishes the cornea and lens
o Produced in the ciliary body by diffusion and active

transport from plasma.

o Flows through the pupil and fills the anterior chamber of

the eye.
o Reabsorbed through a network of trabeculae into the

canal of Schlemm, a venous channel at the junction between the iris and the cornea (anterior chamber angle). Obstruction of this outlet leads to increased intraocular pressure
Ganongs Review of Med Physiol, 23rd ed

Physical Principles of Optics

Refraction of Light Refractive Index of a Transparent Substance Light rays travel through air at a velocity of about 300,000 km/sec, (the refractive index of air is 1.00), but they travel much slower through transparent solids and liquids. The refractive index of a transparent substance is the ratio of the velocity of light in air to the velocity in the substance. If light travels through a particular type of glass at a velocity of 200,000 km/sec, the refractive index of this glass is 300,000 divided by 200,000, or 1.50.

Guyton & Hall Textbook of Med Physiol, 12e, 2011

Physical Principles of Optics

Guyton & Hall Textbook of Med Physiol, 12e, 2011

Figure 49-1 Light rays entering a glass surface perpendicular to the light rays (A) and a glass surface angulated to the light rays (B). This figure demonstrates that the distance between waves after they enter the glass is shortened to about two-thirds that in air. It also shows that light rays striking an angulated glass surface are bent.

Refraction of Light Rays at an Interface Between Two Media with Different Refractive Indices
A. Light rays strike an interface between two media with different refractive indices, perpendicular to the beam rays enter the second medium without deviating, but with decreased velocity of transmission and shorter wavelength B. Light rays pass through an angulated interface, the rays bend if the refractive indices of the two media are different (leaving air, refractive index of 1.00, entering a block of glass, refractive index of 1.50)

Guyton & Hall Textbook of Med Physiol, 12e, 2011

Refraction of Light Rays at an Interface Between Two Media with Different Refractive Indices
Beam enters the lower edge of the glass ahead of the upper edge. The wave front in the upper portion of the beam continues to travel at a velocity of 300,000 km/sec, while that which entered the glass travels at a velocity of 200,000 km/sec the upper portion of the wave front moves ahead of the lower portion so that the wave front is no longer vertical but angulated to the right. Because the direction in which light travels is always perpendicular to the plane of the wave front, the direction of travel of the light beam bends downward. This bending of light rays at an angulated interface is known as refraction. Note that the degree of refraction increases as a function of (1) the ratio of the two refractive indices of the two transparent media (2) the degree of angulation between the interface and the entering wave front.
Guyton & Hall Textbook of Med Physiol, 12e, 2011

Application of Refractive Principles to Lenses

Convex Lens Focuses Light Rays

Figure 49-2 Bending of light rays at each surface of a convex spherical lens, showing that parallel light rays are focused to a focal point.
Guyton & Hall Textbook of Med Physiol, 12e, 2011

Convex Lens Focuses Light Rays

Parallel light rays passing through the center of the convex lens strike the lens exactly perpendicular to the lens surface and, therefore, pass through the lens without being refracted Toward either edge of the lens, the light rays strike a progressively more angulated interface The outer rays bend more toward the center, convergence of the rays. Half the bending occurs when the rays enter the lens, and half as they exit from the opposite side. Parallel light rays passing through each part of the lens will be bent that all the rays will pass through the focal point.
Guyton & Hall Textbook of Med Physiol, 12e, 2011

Application of Refractive Principles to Lenses

Concave Lens Diverges Light Rays

Figure 49-3 Bending of light rays at each surface of a concave spherical lens, showing that parallel light rays are diverged. Guyton & Hall Textbook of Med Physiol, 12e, 2011

Concave Lens Diverges Light Rays

The parallel light rays that enter the center of the concave lens strike interface that is perpendicular to the beam and, therefore, do not refract. The rays at the edge of the lens enter the lens ahead of the rays in the center. This is opposite to the effect in the convex lens, and it causes the peripheral light rays to diverge from the light rays that pass through the center of the lens. Thus, the concave lens diverges light rays, but the convex lens converges light rays.

Guyton & Hall Textbook of Med Physiol, 12e, 2011

Cylindrical Lens Bends Light Rays in Only One Plane-Comparison with Spherical Lenses

Figure 49-4 A Point focus of parallel light rays by a spherical convex lens. B, Line focus of parallel light rays by a cylindrical convex lens.
Guyton & Hall Textbook of Med Physiol, 12e, 2011

The relation of focal length of the lens, distance of the point source of light, and distance of focus
Figure 49-6 The two upper lenses have the same focal length, the light rays entering the top lens are parallel, whereas those entering the middle lens are diverging; the effect of parallel versus diverging rays on the focal distance is shown. The bottom lens has far more refractive power than either of the other two lenses
Guyton & Hall Textbook of Med Physiol, 12e, 2011

The relation of focal length of the lens, distance of the point source of light, and distance of focus

Figure 49-6 The effect of parallel versus diverging rays on the focal distance is shown. The bottom lens has far more refractive power than either of the other two lenses (i.e., has a much shorter focal length), demonstrating that the stronger the lens is (the more refractive power tha lens has), the nearer to the lens the point focus is. The relation of focal length of the lens, distance of the point source of light, and distance of focus is expressed by the following formula: 1 = 1+ 1 f a b f is the focal length of the lens for parallel rays, a is the distance of the point source of light from the lens, b is the distance of focus on the other side of the lens.
Guyton & Hall Textbook of Med Physiol, 12e, 2011

Measurement of the Refractive Power of a Lens-"Diopter"

Figure 49-8 Effect of lens strength on the focal distance

Guyton & Hall Textbook of Med Physiol, 12e, 2011

Measurement of the Refractive Power of a Lens-"Diopter"

The more a lens bends light rays, the greater is its "refractive power, measured in terms of diopters. Figure 49-8: a spherical lens that converges parallel light rays to a focal point 1 meter beyond the lens has a refractive power of +1 diopter,. I If the lens is capable of bending parallel light rays twice as much as a lens with a power of +1 diopter, it is said to have a refractive power of +2 diopters, the light rays come to a focal point 0.5 meter beyond the lens. A lens capable of converging parallel light rays to a focal point only 10 centimeters (0.10 meter) beyond the lens has a refractive power of +10 diopters. T The refractive power of concave lenses cannot be stated in terms of the focal distance beyond the lens because the light rays diverge If a concave lens diverges light rays at the same rate that a 1-diopter convex lens converges them, the concave lens is said to have a dioptric strength of 1. Likewise, if the concave lens diverges light rays as much as a +10-diopter lens converges them, this lens is said to have a strength of -10 diopters.
Guyton & Hall Textbook of Med Physiol, 12e, 2011

Refractive Surfaces of the Eye as a Single LensThe "Reduced" Eye

Figure 49-9 The eye as a camera. The numbers are the refractive indices
Guyton & Hall Textbook of Med Physiol, 12e, 2011

Optics of the Eye The Eye as a Camera

The eye, has a lens system, a variable aperture system (the pupil), and a retina. The lens system of the eye is composed of four refractive interfaces: (1) the interface between air and the anterior surface of the cornea, (2) the interface between the posterior surface of the cornea and the aqueous humor, (3) the interface between the aqueous humor and the anterior surface of the lens of the eye, (4) the interface between the posterior surface of the lens and the vitreous humor. The internal index of air is 1; the cornea, 1.38; the aqueous humor, 1.33; the crystalline lens (on average), 1.40; and the vitreous humor, 1.34.

Guyton & Hall Textbook of Med Physiol, 12e, 2011

Refractive Surfaces of the Eye as a Single LensThe "Reduced" Eye

If all the refractive surfaces of the eye considered to be one single lens, ("reduced eye), it has a total refractive power of 59 diopters when the lens is accommodated for distant vision. + 2/3 of the 59 diopters of refractive power of the eye is provided by the anterior surface of the cornea (not by the eye lens). The refractive index of the cornea is markedly different from that of air, whereas the refractive index of the eye lens is not greatly different from the indices of the aqueous humor and vitreous humor The total refractive power of the internal lens of the eye is only 20 diopters, + 1/3 the total refractive power of the eye. The importance of the internal lens is that, in response to nervous signals from the brain, its curvature can be increased markedly to provide "accommodation,"
Guyton & Hall Textbook of Med Physiol, 12e, 2011

Mechanism of "Accommodation"

Figure 49-10 Mechanism of accommodation (focusing).

Guyton & Hall Textbook of Med Physiol, 12e, 2011

Mechanism of "Accommodation"
In children, and a young person the refractive power of the lens of the eye can be increased voluntarily from 20 diopters to about 34 diopters; "accommodation" of 14 diopters. the shape of the lens is changed from that of a moderately convex lens to that of a very convex lens. the lens is composed of a strong elastic capsule filled with viscous, proteinaceous, transparent fluid. When the lens is in a relaxed state with no tension on its capsule, it assumes an almost spherical shape, owing mainly to the elastic retraction of the lens capsule. However, as shown in Figure 49-10, about 70 suspensory ligaments attach radially around the lens, pulling the lens edges toward the outer circle of the eyeball. These ligaments are constantly tensed by their attachments at the anterior border of the choroid and retina. The tension on the ligaments causes the lens to remain relatively flat under normal conditions of the eye.

Formation of an Image by a Convex Lens

Figure 49-7 A, Two point sources of light focused at two separate points on opposite sides of the lens. B, Formation of an image by a convex spherical lens. Guyton & Hall Textbook of Med Physiol, 12e, 2011

Formation of an Image by a Convex Lens

Figure 49-7A Light rays pass through the center of a convex lens without being refracted in either direction, the light rays from each point source of light come to a point focus on the opposite side of the lens directly in line with the point source and the center of the lens. Figure 49-7B Each point source of light on the object comes to a separate point focus on the opposite side of the lens in line with the lens center. The image of the object, is upside down with respect to the original object, and the two lateral sides of the image are reversed. This is the method by which the lens of a camera focuses images on film.

Measurement of the Refractive Power of a Lens-"Diopter"

The more a lens bends light rays, the greater is its "refractive power. , measured in terms of diopters. The refractive power in diopters of a convex lens is equal to 1 meter divided by its focal length. A spherical lens that converges parallel light rays to a focal point 1 meter beyond the lens has a refractive power of +1 diopter, (Figure 49-8). A lens capable of bending parallel light rays twice as much as a lens with a power of +1 diopter, it is said to have a strength of +2 diopters, and the light rays come to a focal point 0.5 meter beyond the lens. A lens capable of converging parallel light rays to a focal point only 10 centimeters (0.10 meter) beyond the lens has a refractive power of +10 diopters. Concave lenses "neutralize" the refractive power of convex lenses. Thus, placing a 1-diopter concave lens immediately in front of a 1-diopter convex lens results in a lens system with zero refractive power

Errors of Refraction

Figure 49-12 Parallel light rays focus on the retina in emmetropia, behind the retina in hyperopia, and in front of the retina in myopia

Correction of Myopia and Hyperopia by Use of Lenses

Figure 49-13 Correction of myopia with a concave lens, and correction of hyperopia with a convex lens.

Accommodation

FIGURE 4.11

The eye as an optical device. During fixation the center of the image falls on the fovea. A, With the lens flattened, parallel rays from a distant object are brought to a sharp focus. B, Lens curvature increases with accommodation, and rays from a nearby object are focused.

Accommodation

The solid lines represent the shape of the lens, iris, and ciliary body at rest, and the dashed lines represent the shape during accommodation. When gaze is directed at a near object, ciliary muscles contract. This decreases the distance between the edges of the ciliary body and relaxes the lens ligaments, and the lens becomes more convex.

Refractive Errors Corections

Focusing point sources of light.

Common Defects of the Image-Forming Mechanism

Ganongs Review of Med Physiol, 23rd ed

Common defects of the optical system of the eye. In myopia (nearsightedness), the eyeball is too long and light rays focus in front of the retina. Placing a biconcave lens in front of the eye causes the light rays to diverge slightly before striking the eye, so that they are brought to a focus on the retina.

Common Defects of the Image-Forming Mechanism

Ganongs Review of Med Physiol, 23rd ed

Common defects of the optical system of the eye. In hyperopia (farsightedness), the eyeball is too short and light rays come to a focus behind the retina. A biconvex lens corrects this by adding to the refractive power of the lens of the eye.

RETINA
o o

Extends anteriorly almost to the ciliary body Organized in 10 layers and contains the rods and cones, which are the visual receptors, plus four types of neurons: bipolar cells, ganglion cells, horizontal cells, and amacrine cells The rods and cones synapse with bipolar cells, and the bipolar cells synapse with ganglion cells The axons of the ganglion cells converge and leave the eye as the optic nerve Horizontal cells connect receptor cells to the other receptor cells in the outer plexiform layer Amacrine cells connect ganglion cells to one another in the inner plexiform layer via processes of varying length and patterns Gap junctions also connect retinal neurons to one another, and the permeability of these gap junctions is regulated.
Ganongs Review of Med Physiol, 23rd ed

o o o o o

RETINA

The receptor layer of the retina rests on the pigment epithelium next to the choroid light rays must pass through the ganglion cell and bipolar cell layers to reach the rods and cones The pigment epithelium absorbs light rays, preventing the reflection of rays back through the retina. Such reflection would produce blurring of the visual images.

Ganongs Review of Med Physiol, 23rd ed

RETINA

A photograph of the optic fundus

Ganongs Review of Med Physiol, 23rd ed The optic nerve leaves the eye and the retinal blood vessels enter it at a point 3 mm medial to and slightly above the posterior pole of the globe. This region is visible through the ophthalmoscope as the optic disk (Figure 123). There are no visual receptors over the disk, and consequently this spot

RETINA

An illustration of the optic fundus

Ganongs Review of Med Physiol, 23rd ed The optic nerve leaves the eye and the retinal blood vessels enter it at a point 3 mm medial to and slightly above the posterior pole of the globe. This region is visible through the ophthalmoscope as the optic disk (Figure 123). There are no visual receptors over the disk, and consequently this spot

Tahap-tahap persepsi penglihatan

cahaya kornea susunan optik mata penajaman rangsang di retina (fovea sentralis)
di retina cahaya diubah menjadi listrik oleh fotoreseptor (transduksi) potensial reseptor potensial aksi n. optikus (N. II) tr. optikus korteks penglihatan di proses persepsi

RETINA

Neural components of the extrafoveal portion of the retina. C, cone; R, rod; MB (miget bipolara), RB (rod bipolar), and FB (flat bipolar) cells; DG (diffuse ganglion) and MG (midget ganglion) cells; H, horizontal cells; A, amacrine cells. Ganongs Review of Med Physiol, 23rd

Organization of the human retina. A, Choroid. B, Pigment epithelium. C, Photoreceptor layer. D, Neural network layer. E, Ganglion cell layer. r, rod; c, cone; h, horizontal cell; b, bipolar cell; a, amacrine cell; g, ganglion cell. (Modified from Dowling JE, Boycott BB. Organization of the primate retina: Electron microscopy. Proc Roy Soc Lond 1966:166:80111.

Guyton Textbook of Med Physiol 11 th ed

The Retina and Its Photoreceptors

o The retina is a multilayered (10 layers ) structure containing the photoreceptor cells and a complex web of several types of nerve cells o A simpler four-layer scheme: o pigment epithelium, o photoreceptor layer o neural network layer o ganglion cell layer visual signal light entering the retina

The Retina and Its Photoreceptors

o Pigment Epithelium o consists of cells with high melanin content o opaque material o also extends between portions of individual rods and cones

o prevents the scattering of stray light greatly sharpening the resolving power of the retina
o ensures that a tiny spot of light (or a tiny portion of an image) will excite only those receptors on which it falls directly o albinism lack this pigment blurred vision o phagocytose bits of cell membrane that are constantly shed from the outer segments of the photoreceptors.

The Retina and Its Photoreceptors

o Photoreceptor Layer o the rods and cones are packed tightly side-by-side o density of thousands per square millimeter, depending on the region of the retina o each eye contains about 125 million rods and 5.5 million cones o the photoreceptor cells occupy a deep layer of the retina light must pass through several overlying layers to reach them o divided into two classes o the cones are responsible for photopic (daytime) vision, which is in color (chromatic) o the rods are responsible for scotopic (nighttime) vision, which is not in color. o their functions are basically similar o they have structural and biochemical differences.

The Retina and Its Photoreceptors

o Cones Three different photopigments are associated with cone cells that differ in the wavelength of light that optimally excites them. The peak spectral sensitivity for the red-sensitive pigment is 560 nm in red photoreceptors the green-sensitive pigment is 530 nm in green photoreceptors the blue-sensitive pigment, is 420 nm in blue photoreceptors At wavelengths away from the optimum, the pigments still absorb light but with reduced sensitivity Colorblind individuals loss of a single color system produces dichromatic vision lack of two of the systems causes monochromatic vision. all three are lacking, vision is monochromatic and depends only on the rods.

The Retina and Its Photoreceptors

o Rods Long, slender, and cylindrical and is larger than a cone cell.

Its outer segment contains photoreceptor disks composed of cellular membrane in which the molecules of the photopigment rhodopsin are embedded
The lamellae near the tip are regularly shed and replaced with new membrane synthesized at the opposite end of the outer segment. The inner segment, connected to the outer segment by a modified cilium, contains the cell nucleus, many mitochondria that provide energy for the phototransduction process, and other cell organelles. At the base of the cell is a synaptic body that makes contact with one or more bipolar nerve cells and liberates a transmitter substance in response to changing light levels.

Retina
o 2 reseptor batang, kerucut potensial reseptor Batang: scotopic, akromatik, susunan konvergen (receptive field) Kerucut: photopic, kepadatan di fovea & susunan tidak konvergen ( ketajaman penglihatan), penglihatan warna (3 jenis kerucut).

o 4 neuron sel bipolar, sel ganglion, sel horizontal, sel amakrin sel bipolar potensial generator; jawaban berbeda thd warna, intensitas, bentuk, perspekstif dan gerak sel ganglion (N. II) potensial aksi, mengkode citra (oncenter/off-center); menghantarkan rentetan potensial aksi melalui jaras paralel yang ke korteks penglihatan

Visual Pathway
Transection of the pathways at the locations indicated by the letters causes the visual field defects shown in the diagrams on the right. The fibers from the nasal half of each retina decussate in the optic chiasm, so that the fibers in the optic tracts are those from the temporal half of one retina and the nasal half of the other. A lesion that interrupts one optic nerve causes blindness in that eye (A). A lesion in one optic tract causes blindness in half of the visual field (C) and is called homonymous (same side of both visual fields) hemianopia (half-blindness). Lesions affecting the optic chiasm destroy fibers from both nasal hemiretinas and produce a heteronymous (opposite sides of the visual fields) hemianopia (B). Occipital lesions may spare the fibers from the macula (as in D) because of the separation in the brain of these fibers from the others subserving

Pathways to the Cortex

Ganglion cell projections from the right hemiretina of each eye to the right lateral geniculate body and from this nucleus to the right primary visual cortex. Note the six layers of the geniculate. P ganglion cells project to layers 36, and M ganglion cells project to layers 1 and 2. The ipsilateral (I) and contralateral (C) eyes project to alternate layers. Not shown are the interlaminar area cells, which project via a separate component of the P pathway to blobs in the visual cortex.

Neural Pathways
o

The axons of the ganglion cells pass caudally in the optic nerve and optic tract to end in the lateral geniculate body, a part of the thalamus The fibers from each nasal hemiretina decussate in the optic chiasm. In the geniculate body, the fibers from the nasal half of one retina and the temporal half of the other synapse on the cells whose axons form the geniculocalcarine tract. This tract passes to the occipital lobe of the cerebral cortex The primary visual receiving area (primary visual cortex, Brodmann's area 17; also known as V1), is located principally on the sides of the calcarine fissure (Figure 8-5).
Medial view of the human right cerebral hemisphere showing projection of the retina on the occipital cortex around the calcarine fissure.

Ganongs Review of Med Physiol, 21st ed

Neural Pathways
o

Some ganglion cell axons pass from the optic tract to the pretectal region of the midbrain and the superior colliculus, where they form connections that mediate pupillary reflexes and eye movements.

Pupillary Reflexes
o When light is directed into one eye, the pupil constricts (pupillary light reflex) o The pupil of the other eye also constricts (consensual light reflex) o The impulses leave the optic nerves near the lateral geniculate bodies enter the midbrain via the brachium of the superior colliculus terminate in the pretectal nucleus the second-order neurons project to the ipsilateral Edinger-Westphal nucleus and the contralateral Edinger-Westphal nucleus the third-order neurons pass to the ciliary ganglion in the oculomotor nerve the fourth-order neurons pass from this ganglion to the ciliary body. This pathway is dorsal to the pathway for the near response.
Ganongs Review of Med Physiol, 21st ed

Neural Pathways
o

The frontal cortex is also concerned with eye movement, and especially its refinement

o
o o

The bilateral frontal eye fields in this part of the cortex are concerned with control of saccades
An area just anterior to these fields is concerned with vergence and the near response The frontal areas concerned with vision probably project to the nucleus reticularis tegmentalis pontinus, and from there to the other brain stem nuclei mentioned above.

Ganongs Review of Med Physiol, 21st ed

The Receptors

The Receptors
o

Rods and cones are divided into

o

an outer segment
o

made up of regular stacks of flattened saccules or disks containing the photosensitive compounds that react to light, initiating action potentials in the visual pathways rich in mitochondria

an inner segment that includes a nuclear region

o

a synaptic zone

THE PHOTORECEPTOR MECHANISM

Electrical Responses The action of light on photosensitive compounds in the rods and cones light is absorbed structural changes electrical potential changes initiate action potentials in the retina

The receptor potentials of the photoreceptors and the electrical responses of most of the other neural elements in the retina are local, graded potentials Only in the ganglion cells that all-or-none action potentials transmitted over appreciable distances are generated The responses of the rods, cones, and horizontal cells are hyperpolarizing The responses of the bipolar cells are either hyperpolarizing or depolarizing Amacrine cells produce depolarizing potentials and spikes that may act as generator potentials for the propagated spikes produced in the ganglion cells.

Figure 1211
Intracellularly recorded responses of cells in the retina to light. The synaptic connections of the cells are also indicated. The eye is unique in that the receptor potentials of the photoreceptors and the electrical responses of most of the other neural elements in the retina are local, graded potentials. The rod (R) on the left is receiving a light flash, whereas the rod on the right is receiving steady, low-intensity illumination. The responses of rods and horizontal cells (H) are hyperpolarizing, responses of bipolar cells (B) are either hyperpolarizing or depolarizing, and amacrine (A) cells produce depolarizing potentials and spikes that may act as generator potentials for propagated spikes of ganglion cells (G).

Ionic Basis of Photoreceptor Potentials

o

Na+ channels in the outer segments of the rods and cones are open in the dark, so current flows from the inner to the outer segment Current also flows to the synaptic ending of the photoreceptor. The Na+K+ pump in the inner segment maintains ionic equilibrium.

o o

Release of synaptic transmitter is steady in the dark.

When light strikes the outer segment, the reactions that are initiated close some of the Na+ channels, and the result is a hyperpolarizing receptor potential.

The hyperpolarization reduces the release of synaptic transmitter, and this generates a signal in the bipolar cells that ultimately leads to action potentials in ganglion cells.
The action potentials are transmitted to the brain.

Effect of light on current flow in visual receptors. In the dark, Na+ channels in the outer segment are held open by cGMP. Light leads to increased conversion of cGMP to 5'-GMP, and some of the channels close. This produces hyperpolarization of the synaptic terminal of the photoreceptor.

Initial steps in phototransduction in rods. Light activates rhodopsin, which activates transducin to bind GTP. This activates phosphodiesterase, which catalyzes the conversion of cGMP to 5'-GMP. The resulting decrease in the cytoplasmic cGMP concentration causes cGMP-gated ion channels to close.

Figure 819

Sequence of events involved in phototransduction in rods and cones.

EYE MOVEMENTS
A very high order of coordination of the movements of the two eyes is necessary if visual images are to fall at all times on corresponding points in the two retinas and diplopia is to be avoided. o Four types of eye movements, each controlled by a different neural system but sharing the same final common path, the motor neurons that supply the external ocular muscles o Saccades, sudden jerky movements, occur as the gaze shifts from one object to another, programmed in the frontal cortex and the superior colliculi, bring new objects of interest onto the fovea and reduce adaptation in the visual pathway that would occur if gaze were fixed on a single object for long periods. o Smooth pursuit movements are tracking movements of the eyes as they follow moving objects, programmed in the cerebellum o Vestibular movements, adjustments in response to stimuli initiated in the semicircular canals, maintain visual fixation as the head moves. o Convergence movements bring the visual axes toward each other as attention is focused on objects near the observer.
o

EYE MOVEMENTS

Extraocular muscles subserving the six cardinal positions of gaze. The eye is adducted by the medial rectus and abducted by the lateral rectus. The adducted eye is elevated by the inferior oblique and depressed by the superior oblique; the abducted eye is elevated by the superior rectus and depressed by the inferior rectus. (Reproduced, with permission, from Greenberg DA, Aminoff MJ, Simon RP: Clinical Neurology, 5th ed. McGraw-Hill, 2002.)

EYE MOVEMENTS

Figure 8-8. The six extraocular muscles, viewed from the top. (Modified from Dox I, Melloni BJ, Eisner GM: Melloni's Illustrated Medical Dictionary. Williams & Williams, 1979.)

EYE MOVEMENTS

Figure 1222 Figure 1222

Fig 12-22
Extraocular muscles subserving the six cardinal positions of gaze. The eye is adducted by the medial rectus and abducted by the lateral rectus. The adducted eye is elevated by the inferior oblique and depressed by the superior oblique; the abducted eye is elevated by the superior rectus and depressed by the inferior rectus. (From Squire LR, et al [editors]: Fundamental Neuroscience, 3rd ed. Academic Press, 2008.)

Visual Acuity
o

Visual acuity is the degree to which the details and contours of objects are perceived. Visual acuity is usually defined in terms of the minimum separableie, the shortest distance by which two lines can be separated and still be perceived as two lines. Visual threshold (!) is the minimal amount of light that elicits a sensation of light

Visual Acuity
Clinically, is often determined by the use of the Snellen letter charts viewed at a distance of 20 ft (6 m). The individual being tested reads aloud the smallest line distinguishable. The results are expressed as a fraction. The numerator of the fraction is 20, the distance at which the subject reads the chart. The denominator is the greatest distance from the chart at which a normal individual can read the smallest line the subject can read. Normal visual acuity is 20/20; a subject with 20/15 visual acuity has better than normal vision (not farsightedness); and one with 20/100 visual acuity has subnormal vision.

The Snellen charts are designed so that the height of the letters in the smallest line a normal individual can read at 20 ft subtends a visual angle of 5 minutes.
Each of the lines in the letters are separated by 1 minute of arc. Thus, the minimum separable in a normal individual corresponds to a visual angle of about 1 minute.

Visual Fields & Binocular Vision

The visual field of each eye is the portion of the external world visible out of that eye.

Theoretically, it should be circular, but actually it is cut off medially by the nose and superiorly by the roof of the orbit
The peripheral portions of the visual fields are mapped with an instrument called a perimeter, and the process is referred to as perimetry. One eye is covered while the other is fixed on a central point. A small target is moved toward this central point along selected meridians, and, along each, the location where the target first becomes visible is plotted in degrees of arc away from the central point The central visual fields are mapped with a tangent screen, a black felt screen across which a white target is moved. By noting the locations where the target disappears and reappears, the blind spot and any objective scotomas (blind spots due to disease) can be outlined.

Visual Fields & Binocular Vision

Monocular and binocular visual fields. The dashed line encloses the visual field of the left eye; the solid line, that of the right eye. The common area (heart-shaped clear zone in the center) is viewed with binocular vision. The colored areas are viewed with monocular vision.

Binocular Vision

The central parts of the visual fields of the two eyes coincide; therefore, anything in this portion of the field is viewed with binocular vision.

The impulses set up in the two retinas by light rays from an object are fused at the cortical level into a single image (fusion)
The points on the retina on which the image of an object must fall if it is to be seen binocularly as a single object are called corresponding points. If one eye is gently pushed out of the line while staring fixedly at an object in the center of the visual field, double vision (diplopia) results; the image on the retina of the eye that is displaced no longer falls on the corresponding point.

Binocular vision has an important role in the perception of depth. However, depth perception also has numerous monocular components, such as the relative sizes of objects, the degree one looks down at them, their shadows, and, for moving objects, their movement relative to one another (movement parallax).

Dark Adaptation

The decline in visual threshold is known as dark adaptation. It is nearly maximal in about 20 minutes, although some further decline occurs over longer periods.

The dark adaptation response actually has two components

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The first drop in visual threshold, rapid but small in magnitude, is known to be due to dark adaptation of the cones because when only the foveal, rod-free portion of the retina is tested, the decline proceeds no further. In the peripheral portions of the retina, a further drop occurs as a result of adaptation of the rods. The total change in threshold between the light-adapted and the fully dark-adapted eye is very great.

This rise in visual threshold is known as light adaptation. occurs over a period of about 5 minutes. It is merely the disappearance of dark adaptation.

Dark Adaptation

Range of luminance to which the human eye responds, with the receptive mechanisms involved. (Reproduced, with permission, by courtesy of Campbell FW, from Bell GH, Emslie-Smith D, Paterson CR: Textbook of Physiology and Biochemistry, 9th ed. Churchill Livingstone, 1976.)

Dark Adaptation