INTRODUCTORY LECTURE ON THE PHYSIOLOGY OF VISION S. I. OGUNGBEMI DEPARTMENT OF PHYSIOLOGY UNIVERSITY OF LAGOS SPECIAL SENSES Special senses are: 1. Vision 2. Audition 3. Olfaction 4. Gustation and 5. Special proprioception (Vestibular apparatus) They serve as tools for learning and formation of memory Monday, July 31, 2017 2 CHAPTERS OF THE LECTURE 1. Functional Anatomy of the Eye 2. Optics and Optical Defects 3. The Visual Pathway 4. Photoreceptor Mechanism of Retina Monday, July 31, 2017 3 Visual System • Eye • Accessory structures – Eyebrows, eyelids, eyelashes, tear glands – Protect eyes from sunlight and damaging particles • Optic nerve (II) – Tracts – Pathways • Eyes respond to light and initiate afferent action potentials 15-4 Accessory Structures of Eye• • • • • Eyebrows – Prevent running perspiration into eyes – Shade Eyelids or palpebrae – Consist of 5 tissue layers – Protect and lubricate Conjunctiva – Covers inner eyelid and anterior part of eye Lacrimal apparatus Extrinsic eye muscles 15-5 Lacrimal Apparatus • Lacrimal apparatus – Lacrimal Gland: Produces tears to moisten, lubricate, wash • Lacrimal Canaliculi – Collects excess tears • Punctum • Lacrimal Sac • Nasolacrimal duct – Opens into nasal cavity 15-6 Extrinsic Eye Muscles 15-7 Anatomy of the Eye • Three coats or tunics – Fibrous: Consists of sclera and cornea – Vascular: Consists of choroid, ciliary body, iris – Nervous: Consists of retina 15-8 Anatomy of the Eye • Fibrous tunic: Outer – Sclera: White outer layer, maintains shape, protects internal structures, provides muscle attachment point, continuous with cornea – Cornea: Avascular, transparent, allows light to enter eye and bends and refracts light • Vascular tunic: Middle – Iris: Controls light entering pupil; smooth muscle – Ciliary muscles: Control lens shape; smooth muscle • Retina: Inner – Contains neurons sensitive to light – Macula lutea or fovea centralis: Area of greatest visual acuity – Optic disc: Blind spot • Compartments – Anterior: Aqueous humor – Posterior: Vitreous humor • Lens – Held by suspensory ligaments attached to ciliary muscles – Transparent, biconvex 15-9 Horizontal section of the right eye. AP, anterior pole; PP, posterior pole; VA, visual axis Monday, July 31, 2017 10 1. Functional Anatomy of the Eye The eye has 3 concentric coats or layers for the specialised sense of vision. The sclera – the outer protective layer of fibrous coat which is transparent anteriorly as the cornea i.e. ⅙ of sclera The choroid – Middle melanin-pigmented layer which contains the blood vessels which nourish the eye. The specialised anterior portions are ciliary body and iris The retina – the innermost layer which contains the photoreceptor cells i.e. the rods and cones. Monday, July 31, 2017 11 The Sclera • Sclera is the tough posterior outer coat. • It is composed of tightly bound elastic and collagen fibres. • It provides adequate protection for internal contents and components of the eye. • It withstands high intraocular pressure of about 20 mmHg. • The high intraocular pressure keeps structures involved in vision in proper shape and position. Monday, July 31, 2017 12 The Cornea • The cornea is more convex than sclera. • It is the anterior surface of the eyeball. • It is made of collagen fibrils. • It is covered anteriorly by stratified epithelium. • It is continuous with conjunctiva covering the exposed sclera. • It transmits and focuses incident light to the retina. Monday, July 31, 2017 13 • Its epithelium utilises lacrymal secretion to keep cornea in hydrated state. • If the cornea is not hydrated, it dries up and looses its transparency → xerophthalmia. • It has its free nerve endings from trigeminal nerve. • It is avascular – i.e. it has no blood vessel in itself. • Being most, it receives oxygen from its own metabolism directly from atmosphere. Monday, July 31, 2017 14 The Choroid, Ciliary Body and Iris. • Choroid comprises an outer pigmented and inner vascular layer. • Blood vessels of the choroid nourish the inner layers of retina by simple diffusion. • Melanocytes are abundant in the pigmented layer. • Black melanin pigments serve to absorb light rays thereby preventing their reflection back to the retina, which would blur the optical image. Monday, July 31, 2017 15 • Ciliary body arises from the anterior end of the choroid coat. • The ciliary apparatus consists of the ciliary muscles, the iris and the suspensory ligament which suspends the lens. • Ciliary muscle consists of an outer ciliary muscle and inner ciliary processes. • It is made up of radial (dilator) and circular (constrictor) muscles. • Motor supply to the ciliary muscle is parasympathetic with cell bodies of the preganglionic neurons in the EdingerWestphal nucleus of oculomotor nerve CN III Monday, July 31, 2017 16 • Postganglionic neurons are in the ciliary ganglion. • Contraction of ciliary muscle makes the lens more convex. • The iris - thin pigmented contractile diaphragm arising from the ciliary body. • The iris varies the aperture (or pupil) of the lens in response to contraction of the ciliary muscles. • The pupil is the visible coloured aperture of the eye. • The pupil which varies the amount of light entering the eyes. • It contains radial and circular multiunit smooth muscles 17 which control the size of the pupil. Monday, July 31, 2017 • Parasympathetic stimulation causes contraction of the ciliary circular muscles (sphincter pupillae in the iris) and constriction of the pupil. • Sympathetic stimulation causes contraction of radial muscles (dilator pupillae in the iris) and dilation of the pupil. • Sympathetic fibres relay via the superior cervical ganglion. The Aqueous Humour • It is a clear liquid located between cornea and lens • It is continuously formed by active transport and diffusion by the processes of the ciliary body. Monday, July 31, 2017 18 • It nourishes the lens and cornea and also buffers acids produced by the anaerobic glycolysis taking place in the lens. • Its composition is similar to CSF or plasma without proteins. • It is reabsorbed through the Canal of Schlemn into the intrascleral veins at The junction of the iris with the cornea. • Blockage of canal increases aqueous humour volume and intraocular pressure (which is normally 15 – 20 mmHg), leading to glaucoma. Monday, July 31, 2017 19 • Glaucoma can damage the retina and optic nerve and blindness may result. The Lens • It is a biconvex, transparent and elastic solid disc. • It is held in position by suspensory ligament between the iris (in front) and the vitreous humor (behind). • The suspensory ligament attaches it to the posterior surfaces of the ciliary processes. • Reduction of tension in the ligament by contraction of ciliary muscles increases the refractive power of the lens. Monday, July 31, 2017 20 • With advancing age, the lens becomes less elastic resulting in presbyopia. • The function of lens is to provide a fine adjustment to the focus. • No blood vessels in the lens. • The central artery which supplied it before birth atrophies and remains as an end-artery supplying only the retina. The Retina • The retina is inverted. Monday, July 31, 2017 21 • It consists of an outer pigmented layer, that separate it from the choroid, and an inner nervous layer towards the vitreous humour. • The Pigmented layer of the retina and choroid are single sheets of melanin-containing epithelial cells (melanocytes). • The functions of the pigmented layer of the retina are: Absorption of light to prevent reflection blurred image in the eye. Phagocytosis of degenerating membrane discs and shelves from rods and cones. Storage of large quantities of vitamin A which is required for synthesis of visual pigment. Monday, July 31, 2017 22 • Retinal detachment :detachment of pigmented layer from the nervous layer causes blindness. • Treatment → laser surgical attachment of the 2 layers. The Nervous Layer • It comprises rods (for poor light vision) and cones (bright light vision) - outermost, light sensitive and abuts on the pigmented epithelium. • The rods and cones are the visual receptors. – Bipolar cells – middle layer – Ganglion cells – innermost layer Monday, July 31, 2017 23 Monday, July 31, 2017 Schematic diagram of a rod and a cone 24 • In each retina, there are about: • 100 million rods • 7 million cones and • 1 million ganglion cells. • There are also 2 groups of interneurones 1. The horizontal cells which interconnect adjacent rods and cones. 2. The amacrine cells which interconnect the ganglion cells. Monday, July 31, 2017 25 • Interneurones have only dendrites but no axons • The links with other neurones are both presynaptic and postsynaptic. • Only the ganglion cells have axons. • Incident light must therefore pass through layers of cells, axons, and blood vessels before reaching the rods and cones (photoreceptors). Fovea Centralis • It is located in the center of the macula lutea (yellow spot) as a small depression in the eye visual axis. • Fovea centralis is point of highest visual acuity in daylight. Monday, July 31, 2017 26 Neural components of the extrafoveal portion of the retina. C, cone; R, rod; MB, RB, and FB, midget, rod, and flat bipolar cells; DG and MG, diffuse and midget ganglion 27 Monday, July 31, 2017 cells; H, horizontal cells; A, amacrine cells • Nerve fibers and blood vessels which pass to and from the optic disc do not pass over the macula lutea but around it. • Its photoreceptors are cones only. The Optic Disk (Blind Spot) • Ganglion cells converge to form the optic nerve • Optic nerve leaves the eye at the optic disk. • It is situated about 3 mm to the nasal (medial) side of the fovea centralis. Monday, July 31, 2017 28 • It is white in colour because axons are myelinated. • No photoreceptors at optic disk hence blind spot. The Vitreous Humor • It is a transparent embryonic tissue of gelatinous consistency. • It fills the posterior chamber between the lens and retina. Monday, July 31, 2017 29 Rod and cone density along the horizontal meridian through the human retina. A plot of the relative acuity of vision in the various parts of the light-adapted eye would parallel the cone density curve; a similar plot of relative acuity of the dark-adapted eye would parallel the 30 Monday, July 31, 2017 rod density curve. Retina seen through the ophthalmoscope in a normal human. The diagram on the left identifies the landmarks in the photograph on the right. Monday, July 31, 2017 31 Functions of the Complete Eye • Visible light: Portion of electromagnetic spectrum detected by human eye • Refraction: Bending of light • Divergence: Light striking a concave surface • Convergence: Light striking a convex surface • Focal point: Point where light rays converge and cross 15-32 2. OPTICS AND OPTICAL DEFECTS Image Formation on the Retina • Refractory surfaces of the eye from the front are: Anterior and posterior surfaces of the cornea Aqueous humour Anterior surface of the lens Posterior surface of the lens Vitreous humour • When all these refractive surfaces are resolved to a single plane, it forms the principal plane. • It lies about 1.5 mm behind the cornea just in front of the lens. Monday, July 31, 2017 • The diopteric power of the eye is about 60 D. 33 • Near point of the eye is the minimum distance (from the eye) from which an object can be properly focused on the retina. • It is about 10 cm in children, 25 cm in adult, and it increases with age. • The recession of the near point with age (beyond 40 cm due to increase plasticity of the lens) is called presbyopia. • The eye focuses a distant object on the retina. • The image that is formed on the retina is inverted, but is interpreted into the upright position by the brain. • The other defects due to refractive errors of the eye are myopia, hypermetropia and astigmatism Monday, July 31, 2017 34 • One is legally blind when his visual acuity falls below 20/200 feet. • In myopia (i.e. short-sightedness), the image is formed in front of the retina due to long eye balls or increased curvature of the cornea. • This defect is corrected by a concave lens which diverges the in-coming light-rays and so allows the lens to focus the image on the retina. • In hypermetropia (i.e. long-sightedness), the image is formed behind the retina due to short eyeballs or reduced curvature of the cornea. Monday, July 31, 2017 35 • The correction is by convex lens which converges the incident rays and allows the lens of the eye to focus the image on the retina. • Astigmatism is due to irregular curvature of the cornea so that the image is refracted to different foci, causing blurring. • It is corrected by a cylindrical lens. It may be associated with myopia. • Presbyopia (i.e. far-sightedness) is due to old age and has been described earlier. It is due to recession of the near point (i.e.) moving away from the eyes. • Visual acuity is tested by Snellen’s Charts. Monday, July 31, 2017 36 Monday, July 31, 2017 37 Common defects of the optical system of the eye. In hyperopia, 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. In myopia, 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. Monday, July 31, 2017 38 Monday, July 31, 2017 39 Monday, July 31, 2017 40 Other Visual Tests Visual Acuity • In visual acuity, distance vision is tested using Snellen’s chart. • Snellen’s chart is a board on which rows of letters are printed with the larger letters at the top. • The figure that mark each of the rows indicates the distance at which the thickness of the line of the individual letter subtends an angle of 1’’ and the height of the letters 5’’ at the eye. • At that distance, the normal eye conveniently distinguish 41 the letters of that particular row. Monday, July 31, 2017 • The test is read at a distance of 6 m at which the normal eye will read the row of the letters marked with figure 6. • The card is placed in a good white light. • One eye is tested at a time while the other eye is kept covered with an opaque disc in a spectacle frame. • Visual acuity is expressed by a fraction of which numerator represents the test distance in meters i.e. the reciprocal of the visual angle in minute. • The denominator the smallest row of letters which can be read at the distance. Monday, July 31, 2017 42 • Thus normal vision is ⁶⁄₆ viz. 6 is the smallest row of letters that can be read at 6 meters. at which distance. • If the subject has hypermetropia, he will be able to read the same line with and without a convex (or positive) lens. • If the subject has myopia, a negative (or concave) lens will improve his acuity. • Inversion of the Retinal Image • Prick a hole in a piece of black paper, hold it in the left hand about 3 inches from one while the other eye is closed. • Look at the sky through the hole. Monday, July 31, 2017 43 • Hold a pin in your right hand so that its head is close to the eye between the paper and the eye. • It appears to be upside down. • The light coming through the hole in the paper casts a direct shadow of the pin’s head on the retina and this shadow is the same way up as the pin itself. Near Point • Hold an open book in front of the bare eye and bring it nearer just before the point it can no longer be seen clearly. • Measure the distance to the eye. Monday, July 31, 2017 44 Focusing and Accommodation Process • Hold a pencil between one eye and the corner of the room and keep the other eye closed. • Attempt to focus both the corner of the room and pencil at the same time. • We can only focus on one thing at a time. • If an open book is placed about 2 feet from the eyes and the viewer sees it through a screen or fine net held at 6 inches from the eyes. • He can see either the mesh of the screen or net or the letters in the book with clarity, but not at the same time. Monday, July 31, 2017 45 • When the mesh is in sharp focus, the book is blurred but when the letters are in sharp focus, the mesh is blurred. • Accommodation enables man to shift his gaze from near to far objects with ease and greater speed than the finest camera of a most skilled photographer. Retina Vessels • Look at the sky through a pin hole in a piece of black paper held close to the eye. • Close the other eye. • Move the paper up and down and side to side. Monday, July 31, 2017 46 • The shadow of retina vessels will be seen • The group of blood vessels seen depends on the direction of the movement. Blind Spot • Make 2 black circles of ⅛ in diameter and 4 in apart. • Hold up the paper in front of the right eye at arm’s length. • Close the left eye and fix the right eye on the left arm mark and bring the paper slowly to the face. • The right hand mark will disappear (i.e. when focused on blind spot) and then reappear as it is brought nearer (i.e. when focused on the fovea centralis). Monday, July 31, 2017 47 Scheiner’s Experiment • Make 2 pin holes in a card of about 2 mm apart (i.e. less than pupil diameter). • Place the card close to one eye while the other is closed and look through the hole at a distant object, say, across the window. • Hold a pencil with its point about 10 ins in front of the eye such that it comes into the field of vision. • While looking at the distant object, the pencil point appears double. • Carefully, slide along an opaque card to cover only one of the pin holes and then focus on the pencil point. 48 Monday, July 31, 2017 • The window cross-piece will then appear double. • Again slide in a card to obscure the same pin hole as before and notice that the double images disappear. After-Images: after-images are of same shape and size with original stimulus • Look intently at a bright circle of white light for 20-30 s in the dark room. • Turn off the light and look fixedly at a black surface where an after-image of the original stimulus will be seen. • This is positive after-image owing to the lack of second stimulus. Monday, July 31, 2017 49 • Repeat the stimulation and quickly transfer the gaze to the centre of white area larger than the original source of light. • Owing to the second stimulus (the white area), the afterimage will be negative and appear as a dark area on the white ground. • Put a piece of red glass in the illuminated box and transfer the gaze to a white area and the after-image will still be negative and tinged with complementary colour of the first stimulus. • The negative after-image of white is black; of red is green; of blue is yellow, and the longer and stronger the primary stimulus, the stronger will be the after-image. Monday, July 31, 2017 50 Streoscope Vision • Thread a needle, note how you do it and how long it takes. • Close one eye and repeat this experiment without your hand touching each other along visual axis and at right angle to the visual axis. • It is easier done by the 2 eyes. Field of Vision • The subject stands facing the examiner with his back to the light at 2 ft. • Each eye must be examined separately. • The examiner closes his eye opposite the closed eye of the subject. Monday, July 31, 2017 51 • The subject open eye should look fix on the examiner’s open eye while the examiner holds his hand midway between himself and the subject. • The examiner then moves his outstretched forefinger from the periphery towards the centre of the visual field. • The subject is then asked to say when he sees the movement of the finger as the same moment as the examiner, provided they both have normal visual field. • The movements of the hand are repeated in all the different meridian of the field. • Thus, the examiner’s field is compared with the that of the Monday, July 31, 2017 52 subject. • The examiner constantly observe the subject’s eye in case of any wandering from the point of fixation, which should be corrected immediately. • The perimeter is a more accurate tool for measuring visual field. Chromatic Aberration • Look at a bright circle of light through a piece of cobalt glass, which transmit only red and blue rays. • A halo of 1 colour will be seen round the light because, the eye cannot focus at once long and short wavelengths. • The halo will be elliptical in shape if the eye has astigmatism. Monday, July 31, 2017 53 Astigmatism • Cover 1 eye and look at the astigmatic fan with the other eye. • The set of radiating lines on the card is astigmatic fan. • All line should appear black, but if some lines appear grey or blurred, astigmatism is present. • It is easier to say which lines are better seen than which lines appeared blurred. Monday, July 31, 2017 54 Simultaneous and Successive Contrasts • Lay a piece of grey paper on a large piece of blue paper and cover both with tissue paper. • The grey will be tinged with yellow other colour background could be used – simultaneous contrast. • Look fixedly at the centre of a small red square on a grey background; then look quickly at a piece of grey paper. • The after image of the red square will be blue-green – successive contrast. • Other colours can also be used. Monday, July 31, 2017 55 3. The Visual Pathways Visual Field is the view seen by one eye without the movement of the head. • Lateral half of the visual field is called the temporal field • Medial half of the visual field is called the nasal field. • The retinal fibres also are similarly divided into nasal and temporal fibres. • Light from the temporal half of the visual field falls on the nasal half of the retina. • Light from the nasal half of the visual field falls on the temporal half of the retina. Monday, July 31, 2017 56 • Light rays from upper right quadrant of the visual field fall on the lower left quadrant of the retina. • Light from the center of the visual field falls on the macula lutea (fovea centralis). • In this way, there is a total inversion of the image formed at the retina. Field of Vision • Place field of vision disk it against your forehead and stare intently, straight ahead, at the target on the disk. • Have your partner move the mobile target, first to one side, then to the other while you continue to stare straight ahead. Monday, July 31, 2017 57 • Record the exact angle at which the mobile target completely disappears from your field of vision on the Data/Analysis Sheet. Functional Structure of the Visual Pathways • Fibers from the nasal half of each retina decussate at the optic chiasma and run in the contralateral optic tract. • Fibers from the temporal half of each retina (rays from the nasal field) run in the ipsilateral optic tract. • Fibers of each optic tract synapse in the lateral geniculate body of the thalamus. Monday, July 31, 2017 58 • Geniculocalcarine fibers pass by way of optic radiations (geniculocalcarine tract) to the primary visual cortex calcarine sulcus occipital lobe (Broadman’s area 17, also known as V1). 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 3–6, 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 59 Monday, July 31, 2017 component of the P pathway to blobs in the visual cortex. Lesions of the Optic Pathway i. Optic nerve lesion results in total blindness in the affected eye (monocular blindness). ii. Optic tract lesion interrupts crossed nasal fibres from the opposite eye and temporal fibres from the same side. • The result is loss of visual field in the temporal side of the opposite eye (nasal fibres) and nasal field of the same side (temporal fibres) resulting in Homonymous hemianopia. • If the right or left optic tract is damaged, it results in left or right homonymous hemianopia respectively. • Homonymous hemianopia refers to the side of the body viz. left- or right-sided visual sides of the body. Monday, July 31, 2017 60 iii. Optic chiasma lesion interrupts fibres from nasal part of both retinae and hence both temporal fields resulting in bitemporal hemianopia. • Bitemporal hemianopia in optic chiasma lesions is an example of heteronymous hemianopia since both temporal fields are affected i.e. the left temporal field is left of the body and the right temporal field is the right side of the body i.e. left and right sided or heteronymous. iv. Optic radiation – results in homonymous hemianopia but the macula fibres are not affected. The macular fibres run separately from the peripheral visual fields and they also have large number of fibres and so are not easily affected by occipital lesions. 61 Monday, July 31, 2017 Visual pathways. Transection of the pathways at the locations indicated by the letters causes the visual field defects shown in the diagrams on the right (see text). Occipital lesions may spare the fibers from the macula (as in D) because of the separation in the 62 brain of these fibers from the others subserving vision . Monday, July 31, 2017 Visual Pathways 15-63 Eye Disorders • Myopia: Nearsightedness – Focal point too near lens, image focused in front of retina • Hyperopia: Farsightedness – Image focused behind retina • Presbyopia – Degeneration of accommodation, corrected by reading glasses • Astigmatism: Cornea or lens not uniformly curved • Strabismus: Lack of parallelism of light paths through eyes • Retinal detachment – Can result in complete blindness • Glaucoma – Increased intraocular pressure by aqueous humor buildup • Cataract – Clouding of lens • Macular degeneration – Common in older people, loss in acute vision • Diabetes – Dysfunction of peripheral circulation 15-64 Connections of the Optic Tract or Visual fibres i. As above to the lateral geniculate body and then to the calcarine/visual cortex for vision. ii. Pass from the optic chiasma to the suprachiasmatic nucleus of the hypothalamus for control of circadian rhythm. iii. From the lateral geniculate nucleus to the pretectal nucleus of the midbrain which then relays through the occulomotor nerve and Edinger-Westphal (pre-tectal) nucleus to mediate the pupillary and consensual light reflexes. Monday, July 31, 2017 65 iv. From the lateral geniculate nucleus to the superior colliculus which then relays to the visual cortex and medial longitudinal bundle to cause eye movements synchronous with that of the body during postural adjustments. v. Into the ventral lateral geniculate nucleus of the thalamus, to help control some of the body’s behavioral functions. Monday, July 31, 2017 66 Responses in the Visual Pathways • 2 types of ganglion cells are found in the retina • Large ganglion cells (magno or M cells) are concerned with movement and sterropsis • Small ganglion cells (parvo or P cells) are concerned with color, texture and shape. • Axons of ganglion cells (optic nerve) project a detailed spatial representation of retina on the lateral geniculate body. Monday, July 31, 2017 67 • Each geniculate body or nucleus contains 6 well defined layers • Layers 1 and 2 have large cells and are called magnocellular • Layers 3-6 have small cells and are called parvocellular • On each side, layers 1, 4 & 6 receive input from the contralateral eye • Layers 2, 3 & 5 receive input from the ipsilateral eye Monday, July 31, 2017 68 • In each layer there is a precise point–to–point representation of the retina. • Only 10-20% of inputs to the lateral geniculate nucleus comes from the retina. • Major inputs occur from the visual cortex and other brain regions. Monday, July 31, 2017 69 Ganglion cells P M Lateral geniculate MagnoParvo-cellular Inter- Via cellular laminar dendrites laminae laminae (layers 3-6) region (layers 1 & 2) Visual Cortex Superficial layer 4C Blobs Deep layer 4C Function Movement, depth, flicker Color vision Color, texture, shape, fine detail Monday, July 31, 2017 ORGANISATION OF THE VISUAL PATHWAYS 70 The Visual Cortex • The Primary Visual Cortex • It is also called Broadman’s area 17 or VI • It receives point-to-point representation from the lateral geniculate body • It processes input in terms of orientation, edges etc Monday, July 31, 2017 71 Medial view of the human right cerebral hemisphere showing projection of the retina on the occipital cortex around the calcarine fissure. Monday, July 31, 2017 72 Other Cortical Areas Concerned with Vision • The primary visual cortex projects to several other areas of the occipital cortex and the brain. • These areas are as listed below. 1. V2,V3, VP – Continued processing, larger visual fields 2. V3A – Motion 3. V4v – Unknown 4. MT/V5 – Motion, control of movement 5. LO – Recognition of objects 6. V7 – Unknown 7. V8 – Color vision Monday, July 31, 2017 73 Some of the main areas to which the primary visual cortex (V1) projects in the human brain. Lateral and medial views Monday, July 31, 2017 74 Visual Reflexes i. Pupillary light reflex ii. Consensual light reflex iii. Accommodation reflex i. Pupillary (Direct) light reflex • When bright light shines into one eye, it results in pupillaryconstriction in the affected eye. • Pathway: retinal cells → pretectal nucleus → Edinger-Westiphal nucleus → occulomotor nerves → pupillary constriction. Monday, July 31, 2017 75 ii. Consensual light reflex • When bright light is shines into one eye, it results in pupillary constriction in both eyes. • Pathway: retinal cells → Edinger-Westiphal nucleus → occulomotor nerves (of both eyes) → pupillary constriction of both eyes. • The pupils of both eyes are constricted by the contraction of their pupillary circular muscle (sphincter pupillae). Monday, July 31, 2017 76 iii. Accommodation Reflex • Accommodation reflex is the reaction of both eyes when they are focused on a near object. • It consists of: i. Pupillary constriction ii. The contraction of the ciliary muscles in order to relax the pull on the suspensory ligament, increasing the curvature of the lens, thus allowing focusing on the retina. iii. The convergence of the eyeballs in order to enable same part of the object to be focused simultaneously on same part of both retinae. Monday, July 31, 2017 77 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. 78 Monday, July 31, 2017 Focus and Accommodation • Emmetropia: Normal resting condition of lens • Far vision: 20 feet or more from eye • Near vision: Closer than 20 feet – Accommodation – Pupil constriction – Convergence15-79 Decline in the amplitude of accommodation in humans with advancing age. The different symbols Monday, July 31, 2017 80 identify data from different studies The Retina • Provides black backdrop for increasing visual acuity • Sensory retina and pigmented retina • Photoreceptors – Rods: Noncolor vision • Rhodopsin reduction: Light adaptation • Rhodopsin production: Dark adaptation – Cones: Color vision 15-81 Photoreceptive Mechanism of the Eye • Cones are for daylight and colour vision. • They have high threshold and are therefore used for bright light. • Rods are for monochromatic and twilight or night vision. • They have low threshold (scotopic vision). • The visual pigment in the rods in rhodopsin derived from the proteins - opsin and retinal, the aldehyde form of vitamin A or retinol. • Light rays cause decomposition of rhodopsin leading to the development of an action potential. Monday, July 31, 2017 82 Rhodopsin Cycle 15-83 Range of luminance to which the human eye responds, with the receptive mechanisms involved.84 Monday, July 31, 2017 • The pigment for cones is not well-known. • The cone pigments are for red sensitivity (erythrolabe), green sensitivity (chlorolabe) and blue sensitivity (cyanolabe). • Rhodopsin = Scotopsin (an Opsin) and retinal. • Retinal exists as cis-retinal in the dark. • When light falls on the retinal cell, cis-retinal is converted to all-trans-retinal. Monday, July 31, 2017 85 • This leads to the formation of intermediates including metarhodopsin II, which then initiates the development of an action potential in the photoreceptor cell via closure of Na+ channels hyperpolarisation • There is decreased transmitter release and excitation of bipolar and ganglion cells causing impulse transmission to the brain. • This mechanism involves cyclic GMP gated ion channels. • Deficiency of vitamin A in the diet will result in nightblindness (nyctalopia) since it is a component of rhodopsin. Monday, July 31, 2017 86 Sensory Receptor Cells 15-87 Rod Cell Hyperpolarization 15-88 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 hyperpolarisation of the synaptic terminal of the photoreceptor. 89 Monday, July 31, 2017 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. Monday, July 31, 2017 90 Sequence of events involved in phototransduction in rods and cones Monday, July 31, 2017 91 Colour Vision • Young-Helmboltz or trichromatic theory proposes three types of cones response to light of 3 different wavelengths/colours i.e. 450 nm (Blue), 550 nm (Green) and 600 nm (Red). • The colour perceived is a combination of the degree to which each of these cones or primary colours are stimulated. • Equal stimulation of the three cones leads to a white colour. Monday, July 31, 2017 92 Absorption spectra of the three cone pigments in the human retina. The S pigment that peaks at 440 nm senses blue, and the M pigment that peaks at 535 nm senses green. The remaining L pigment peaks in the yellow portion of the spectrum, at 565 nm, but its spectrum extends far enough into the long wavelengths to sense red 93 Monday, July 31, 2017 • Colour blindness can occur when any or all (total colour blindness) of the three cone types are absent. • These are dichromats and monochromats. • Trichromats have all the 3 cones and so have normal colour vision. • Monochromats are completely colour blind. • They only have one cone type and see only black and white or grey. Monday, July 31, 2017 94 • Dichromats have two cone types losing either that to Red (protanopia) or Green (deuteranopia). • They confuse red or green and perceive colours in shade of yellow and blue. • Blue cone loss (tritanopia) is rare. • Colour blindness is a sex-linked recessive gene and therefore is more common in males. • Colour blindness is tested by Ishihara charts. Monday, July 31, 2017 95 Monday, July 31, 2017 96 Dark Adaptation • When the eye is exposed to poor light it adapts and becomes increasingly sensitive to the dark resulting in increased visual acuity. • There are two components: An initial rapid adaptation within 5 minutes though with poor detail. • This is accomplished by the cones. Monday, July 31, 2017 97 Then follows a more prolonged adaptation which is about 60% of the total adaptation. • This results to an increase in acuity and is due to the Rods. • Hence the rods contribute a quantitatively greater proportion to visual adaptations in the dark, due to their higher content of photopigment. • Monday, July 31, 2017 98 Dark adaptation. The curve shows the change in the intensity of a stimulus necessary to just excite the retina in dim light as a function of the time the 99 Monday, July 31, 2017 observer has been in the dark. EFFECTS OF AGING ON THE SPECIAL SENSES • Slight loss in ability to detect odors • Decreased sense of taste • Lenses of eyes lose flexibility • Development of cataracts, macular degeneration, glaucoma, diabetic retinopathy • Decline in visual acuity and color perception 15-100
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