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PHYSIOLOGY OF THE EYE AND VISUAL TRANSDUCTION Eye - Complex structure - Spherical with a diameter of approximately 1 inch - Only

the anterior 1/6th of the eye surface is visible - Rest enclosed and protected by a cushion of fat and the wall of the bony orbit - Fat pad occupies most of the orbit Vision - Dominant sense - 70% of all sensory receptors are in the eyes - Nearly half of the cerebral cortex is involved in the processing of visual information - Visual receptor cells (photoreceptors) sense and encode patterns of light that enter the eye - Brain uses these signals to give us images Lacrimal Apparatus - Lacrimal gland - Located superior and lateral to the eye - Continuously releases a dilute saline solution into the superior part of conjunctival sac - Ducts - Drain the excess lacrimal secretions into the nasal cavity Function of Tears - enhanced tearing during eye irritation - to wash away or dilute the irritating substance - enhanced tearing during emotional upset Movement of Tears - through the lacrimal apparatus - tears are released through excretory ducts - blinking spreads the tears downward and across the eyeball to the medial commissure where they enter the paired lacrimal canals (canaliculi) via two tiny openings called lacrimal puncta - from the canals, the tears drain into the lacrimal sac and then into the nasolacrimal duct which empties into the nasal cavity at the inferior nasal meatus Conjuctiva - transparent mucous membrane

- lines the eyelids - Functions Major - produce a lubricating mucus - prevents the eyes from drying out Other - Protection - Prevernts foreign objects from penetrating beyond the confines of its sac Wall of Eyeball - consists of tunics Outermost - Fibrous tunic - sclera - Cornea Middle - Vascular tunic - Ethoroid - Ciliary Body - Iris Innermost - Sensory tunic - retina Outermost Coat (Fibrous Tunic) - consists of the two regions - sclera & cornea - sclera - forms the posterior portion and the bulk of the coat - glistening white and opaque - seen anteriorly as the white of the eye - tough and hard - protects and shapes the eyeball and provides a sturdy anchoring site for the extrinsic eye muscles - continuous with the dura mater posteriorly where it is pierced by the optic nerve - Cornea - allows light to enter the eye - covered with epithelial sheets that help protect the cornea from abrasion - capable of regeneration and repair - well-supplied with nerve endings (mostly pain receptors) - only tissue in the body that can be transplanted from one person to another with little or no possibility of rejection - has no blood vessels - beyond the reach of the immune system

Middle coat (tunic) - pigmented vascular coat - also called the uvea - consists of three regions - Choroid - Ciliary body - Iris - Choroid - highly-vascular dark brown membrane - blood vessels provide nutrients to the entire eye - prevents light scattering within the eye - anteriorly - it becomes the ciliary body - posteriorly - incomplete where the optic nerve leaves the eye - Ciliary Body - consists primarily of interlacing smooth muscle bundles - ciliary muscles control the lens shape - contains folds (ciliary processes) posteriorly that contain capillaries - secretes the fluid that fills the cavity of the anterior segment - suspensory ligament (zonule) extends from the ciliary processes to the lens - helps hold the lens in an upright position Accommodation - Is the ability of the eye to keep an image focused on the retina though the distance between eye and object varies. - Results from contraction of ciliary muscle, which changes the thickness of the lens - To focus on objects closer than 7 meters.

Middle coat (tunic) - Iris - Different colors - contains only one pigment (brown) - presence of a lot of pigment - Eye appear brown of black - Presence of a small amount of pigment - Restricted to the posterior surface - Eye appear blue, green or gray - Scattering of light on the unpigmented parts - New born babies eyes are slate gray - Iris pigment is not yet developed - In close vision and bright light - contraction of circular muscles (pupil constriction) - parasympathetic effect - In distant vision and dim light - contraction of radial muscles (pupil dilation) sympathetic effect

- Contraction of circular or sphincter muscles (miosis), innervated by postganglionic parasympathetic fibers from ciliary ganglion. - Contraction of radial muslces (mydriasis) innervated by postganglionic sympathetic fibers from superior cervical ganglion. Pupillary Light Reflex - Regulation of pupillary size by ambient light levels Fxn: Regulate amount of light entering the eye Quality of retinal image greater depth of focus - Direct light response light directed to one eye causes contriction. - Consensual light response identical constriction in the other eye w/out light stimulation. Inner Coat (Tunic) - delicate, two-layered retina - outer, pigmented layer - inner, transparent neural layer The retina as observed ophthalmoscope - Fovea Centralis with an

- outer, pigmented layer - pigmented epithelial cells - absorb light - prevent it from scattering in the eye - act as phagocytes - store vitamin A - needed by the photoreceptor cells - inner, transparent neural layer - only this layer plays a direct role in vision - composed of 5 types of neurons - either transduce light energy or process light stimuli 1.) photoreceptors (rods and cones) 2.) bipolar cells 3.) ganglion cells 4.) horizontal 5.) amacrine Photoreceptors - modified neurons - Hyperpolarize in response to light - rods (scotopic condition) - More numerous - dim light and peripheral vision receptors - more sensitive to light - do not provide sharp images or color vision - in dim light colors are indistinct - Cones (photopic condition) - operate in bright light - provide high acuity color vision Physiology of Vision - Three types of cones - Red - Blue - Green - Each cone type responds maximally (more strongly) to one color of light - Most light stimulates more than one cone type - Allows us to see a full range of colors Ganglion Cells - ganglion cell axons form the optic nerve - exit via the optic disc

The only part of the visual field that is seen clearly is the tiny part (about 1%) that falls on the fovea centralis. Rapid eye movements shift different parts of the visual field onto the fovea centralis.

Optic Disk

Optic nerve fibers leave the eyeball at the optic disc to form the optic nerve The optic disk is also known as the blind spot

- optic disc ~ blind spot - weak spot in the posterior wall - not reinforced by the sclera - lacks photoreceptors - light focused on it cannot be seen - the brain utilizes a process called filling in so we do not realize gaps in our vision Internal Chambers and Fluids - AQUEOUS HUMOR (clear fluid) - composition ~ blood plasma - forms and drains continually - supplies nutrients and O2 to lens, cornea, retinal cells - removes metabolic wastes - helps maintain IOP - supports the eyeball - VITREOUS HUMOR (clear gel) - forms in embryo - lasts a lifetime - fine collagen fibrils in viscous ground substance - transmits light - supports posterior surface - holds the neural retina firmly against the pigmented layer - helps maintain IOP

Refraction - Speed is constant - when traveling in a given medium - speed changes - when passing from one medium to different one - less dense medium speeds up - more dense medium - slows down Refraction of Light Rays - convex lens surface is thicker in the center - light rays bend - converge or intersect at a single point (focal point) - image formed - real image - upside down - reversed from left to right 2 Major Components of Vertebrate Eye 1. Optical Part 2. Neural Part Refraction Depends on: 1. Radius of curvature 2. Difference in refractive index of two adjoining media. Index of Refraction - A measure of the speed of light within it Refraction of Light Rays - refraction of light rays - three times - moves sequentially from: - air to cornea to aqueous humor to lens to vitreous humor to entire thickness of neural retinal layer to excite photoreceptor cells - light falls on retina: - as an upside-down, left- right-reversed image - brain interprets image as: - right-side-up, correctly oriented left to right

Errors of Refraction 1. Myopia Myopia Nearsightedness Eyes too long Hyperopia Farsightedness Eyes too short

Concave lenses Convex lenses

Astigmatism Diffuse focus Presbyopia 2. 3. 4. Age-related

Uneven cornea or lens Uneven lenses Hardening of lens Convex lenses Generation and Transmission of a Visual Message - light passes through the retina - passes two layers of neurons - ganglion and bipolar cells - light stimulates the photoreceptors (rods & cones) - located near the choroid - absorb light - light-sensitive pigments within the photoreceptors change shape - shape change initiates a series of chemical reactions - result in the generation of an action potential - the impulse travels from the rods and cones to the bipolar neurons and then through the ganglion neurons - ganglion neurons conduct the impulse to the brain via the optic nerve (make right angle turns) Retina = 5 Types of Neurons - Photoreceptors rod and cones - Ganglion Cell - Bipolar interneuron w/in the retina / radially - Horizontal interneuron spread horizontally - Amacrine Phototransduction - A cascade of chemical and electrical event to detect, amplify and to signal a light response. Hyperpolarization - Essential step in relaying the visual signal - Directly modulates the rate of transmitter release from the photoreceptor unto its postsynaptic neurons. Dark Current - Na+ - k+

Hyperopia / Hypermetropia Astigmatism Presbyopia

Visual Acuity - Sharpness of vision. - Depends upon resolving power: - Ability of the visual system to resolve 2 closely spaced dots. - Visual abnormalities - Myopia (nearsightedness): - Hyperopia (farsightedness): - Astigmatism (uneven cornea or lens) - Presbyopia (hardening of the lens- accommodation is impeded)

- Ca 2+ Rhodopsin - G protein = coupled receptor for light - Density =30,000 molecules/sq. micrometer in disk membranes - Rod contains = rhodopsin molecules

RHODOPSIN: 2 Key Components 1. Retinal or Retinol = aldehyde of vitamins A (-500 Da) 2. Opsin = single polypeptide (-41 kDa) - Retinal absorption of photons transduction - II-cis retinal (tail) = kinked & unstable retinal tail - II-cis retinal photons absorption all-trans retinal isomerization (straighter & more stable) - Transducin - G protein - Transduces the light activated signal from rhodopsin into photoreceptors membranes response Transducin (activation by metarhodopsin) losses GDP but gains GTP Stimulation of phosphodiesterase hydrolyzation of cGMP 5- guanylate monophosphate DARK activation of guanyl cyclase GTP Synthesis of cGMP (high in photoreceptor cytoplasm) LIGHT

Stimulation of phosphodiesterase Decrease cGMP Reduce dark current Hyperpolarization of photoreceptors Decrease transmitter release Visual signal to retinal neurons GTP Synthesis of cGMP (high in photoreceptor cytoplasm) Phototransduction Pathway - 1 photon = activates 1 metarhodopsin - transducin activate 700 molecules w/in100 ms - activate phosphodiesterase - Increase cGMP hydrolysis by 100 fold - 1 photon = hydrolysis of 1,400 cGMP - Decrease cGMP by 8% closes 230 of 11,000 cGMP gated channels - Dark current falls by 2%

Dark Adaption 1. 10 minutes = cones 2. 30 minutes = rods - A fully dark-adapted retina, relying on rods have light threshold 15,000 times lower than a retina relying on cones Dark Adaption 2 Mechanisms 1. Neural mechanism - Relatively fast - Low light levels - Multiple mechanisms w/in neuronal network of retina 2. Photoreceptor mechanism - Light rods ineffective (rhodopsin inactivated or bleached) - Dark rods regenerate rhodopsin & become sensitive

- Opsin - couples to transducin - Rhodopsin Kinase - phosphorylates light activated rhodopsin - recognized by arrestin - Arrestin - abundant cytosolic protein - binds to the phosphorylated light-activated rhodopsin - helps terminate the activated state of the receptor

Spectral Sensitivity of Light - Adapted eye - Depends on the photopigments in the cones - Peak of absorbance curves fall at: - 420 nm (violet) - 530 nm (yellow-green) - 560 nm (yellow-red) - 3 cones - Blue short wavelength (S) - Green medium wavelength (M) - Red Long wavelength (L)

Physiology of Vision - Objects have color - they absorb some wavelengths - they reflect some wavelengths - things that look white reflect all wavelengths - black objects absorb all wavelengths - a red apple reflects mostly red light - grass reflects more of the green Color Blindness - congenital lack of cone types - one or more - ex-linked disease (x-linked recessive mutation) - more common in males - 8-10 % - 1 % in females - most common type - red-green - deficit or absence of red or green cones seen as same color - either red or green Inherited defects in Color Vision - X-linked recessive mutation of visual pigment genes - 3 Types 1. Monochromacy - lack of 2 of 3 functional cone pigments - Univariance - S-cone monochromacy ( < .001 % of population) 2. Dichromacy - Absence of one functional pigment - Red: protanopia - Green: deuteranopia - Blue: tritanopia 3. Anomalous trichromacy - Absorption spectrum of one pigment shifted relative to normal - Inability to distinguish certain colors - Red: protanomaly - Green: deuteranomaly - Blue: tritanomaly

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