The Physiology of Vision

Objectives

Review anatomy and histology of light receptors Compare and contrast the rods and Cones Describe the photochemistry of vision Describe the receptor potential of rods and cones Describe dark and light adaptation Explain color vision and color blindness List the cells of the neural pathway Follow the neural pathway up to the visual cortex

Resources: Guytons Textbook of Physiology

Retina

Light sensitive portion of the eye Contains cones for color vision Contains rods for night vision Contains neural architecture Light must pass through the neural elements to strike the light sensitive rods and cones

(1) pigmented layer, (2) layer of rods and cones projecting to the pigment, (3) outer nuclear layer containing the cell bodies of the rods and cones, (4) outer plexiform layer, (5) inner nuclear layer, (6) inner plexiform layer, (7) ganglionic layer, (8) layer of optic nerve fibers, and (9) inner limiting membrane.

The Fovea

A small area at the center of the retina about 1 sq millimeter The center of this area, the central fovea, contains only cones

These cones have a special structure Aid in detecting detail

In the central fovea the neuronal cells and blood vessels are displaced to each side so that the light can strike the cones directly. This is the area of greatest visual acuity

Fovea
Minute area(1sq. mm) in the centre of retina. 100% cones capable of detecting acute and detailed vision. Also has blood vessels, ganglion cells, INL & plexiform layer displaced to side. Light passes un impeded for detailed vision.

Pigment Layer of Retina

Pigment layer of the retina is very important Contains the black pigment melanin Prevents light reflection in the globe of the eye Without the pigment there would be diffuse scattering of light rather than the normal contrast between dark and light. This is what happens in albinos poor visual acuity because of the scattering of light

photoreceptors
There are 2 types of photoreceptors in the retina: rods and cones.

The rods are most sensitive to light and dark changes, shape and movement and contain only one type of light-sensitive pigment. Rods are more numerous than cones in the periphery of the retina. 120 million rods in the human retina.

Cones
The cones are not as sensitive to light as the rods. Cones are most sensitive to one of three different colors (green, red or blue). You cannot see color very well in dark places as cones work in bright light.

Functional segments of rods and cones

Outer segment: Has light sensitive photopigment. Large no.(1000) of discs which are folds of cell membrane. Photopigment is present as transmembrane proteins & form 40% of outer segment mass.

Inner segment: Contains cytoplasm and organelles. Most important mitochondria which provide energy. Nucleus: Synaptic body: Connects to subsequent horizontal and amacrine cells.

Rods

Cones

High sensitivity; specialized for night vision More photopigment High amplification; single photon detection Slow response, long integration time More sensitive to scattered light

Lower sensitivity; specialized for day vision Less photopigment Less amplification Fast response, short integration time More sensitive to direct axial rays

Rods

Cones

low acuity; highly convergent retinal pathways, not present in central fovea achromatic; one type of rod pigment

high acuity; less convergent retinal pathways, concentrated in central fovea chromatic; three types of cones, each with a different pigment that is sensitive to a different part of the visible spectrum (Red, Green and Blue)

e of Rods Cones

Structure of the Rods and Cones

Retinal Rhodopsin visual cycle

Both rods & cones have chemicals which on exposure to light decompose and excite nerve fibres leading from the eye. In rods this light sensitive pigment is RHODOPSIN. In cones, it is colour/ cone pigment that may be; Erythrolabe ( red sensitive) Chlorolabe ( green sensitive) Cyanolabe ( blue sensitive)

Visual cycle
Rhodopsin or visual purple present in the outer segment of retina. It is a combination of protein scotopsin & carotenoid pigment Retinal/ Retinene. Retinal is 11-cis retinal type which is the only form that binds with scotopsin to produce rhodopsin.

Rhodopsin Activation
Light falling on rhodopsin cause it to decompose in a fraction of a second & change from cis form to trans form ( same chemical structure but different conformational structure). The immediate product is Bathorhodopsin that changes into Lumirhodopsin. In microseconds it decays into Metarhodopsin I

In milliseconds it changes to Metarhodopsin II and later more slowly into scotopsin and all trans retinal. Metarhodopsin II is called activated Rhodopsin as it causes electrical changes in the rods that transmit visual signals to the CNS.

Vitamin A & its role in rhodopsin formation

All trans retinal can be converted to 11-cis retinal by an alternate route. This is to convert all trans retinal to all transretinol which is a form of vitamin A. Then all trans retinol is changed to 11-cis retinol under influence of isomerase. Finally 11-cis retinol is converted to 11cisretinal.

Re- formation of Rhodopsin

All trans retinal is converted to 11-cis retinal. This requires energy & enzyme Retinal Isomerase. 11-cis retinal automatically combines with scotopsin to re-form rhodopsin a process occuring in darkness.

Nyctalopia/ Night Blindness

This occurs in severe vitamin A deficiency. Amount of retinal and subsequent rhodopsin are severely decreased. It is called night blindness as the amount of light at night is too little to allow adequate vision in vitamin A deficient people. Person must be on vit. A deficient diet for months to develop nyctalopia as large amounts stored in liver. It can be corrected by a single injection of vit. A in less than an hour.

Photochemistry of Vision

Rod receptor potential

There is hyper polarization rather than depolarization. There is increased negativity of intra rod membrane potential. When rhodopsin decomposes it decreases the rod membrane conductance for Na+ ions in the outer segment of rod.

Inner segment continually pumps Na+ from inside to the outside of rod so ve potential. Outer segment is very leaky to Na+ in dark state. So much of +ve Na leaks back into inside of rod to neutralize most of the negativity.

So in darkness there is decreased negativity of -40 mV rather than -70-80 mV in sensory receptors. When rhodopsin in outer segment is exposed to light & decomposes, it decreases the outer segment membrane conduction for Na+

But the inner segment continues to pump out Na+. So more Na+ leaks out than move back in. There is increased negativity inside rod membrane. Greater the light falls on rods greater is the intra rod membrane negativity hyper polarization.( -70-80 mV)

Sequence of events in Phototransduction

Incident light( photon of light activates electron) Structural change in retinene photo pigment. Conformational change in photo pigment. Activation of G protein Transducin. Activated transducin activates Phosphodiestrase.

Decreased intracellular cGMP. cGMP is destroyed by hydrolysis, before which it was bound with Na channel protin of outer segment to splint it in open state. Closure of sodium channel. Hyper polarization. Decreased release of synaptic transmitter. Response in bipolar & other neural elements Rhodopsin kinase (in a sec) inactivates activated rhodopsin and entire cascade reverses back to normal.

Duration and Sensitivity of the Receptor Potential

A single pulse of light causes activation of the rod receptor potential for more than a second. In the cones these changes occur 4 X faster. Receptor potential is proportional to the logarithm of the light intensity.
very

important for discrimination of the light intensity

Dark and Light Adaptation

In light conditions 1. most of the rhodopsin has been reduced to retinal. 2. much of the retinal of both the rods and the cones will have been converted into vitamin A. Because of these two effects, the concentrations of the photosensitive chemicals remaining in the rods and cones are considerably reduced, and the sensitivity of the eye to light is correspondingly reduced. This is called light adaptationt the eye

In dark conditions
1retinal is converted back to rhodopsin. 2.vitamin A is converted back into retinal to increase light-sensitive pigments,( the final limit being determined by the amount of opsins in the rods and cones to combine with the retinal). This is called dark adaptation Opening and closing of the pupil also contributes to adaptation because it can adjust the amount entering light.

the sensitivity of the retina is very low on first entering the darkness, but within 1 minute, the sensitivity increased 10-fold(that is, the retina can respond to light of one tenth the previously required intensity.) At the end of 20 minutes, the sensitivity has increased about 6000-fold, and at the end of 40 minutes, about 25,000-fold.

Other Mechanisms of Light and Dark Adaptation

change in pupillary size, This can cause adaptation of approximately 30-fold within a fraction of a second because of changes in the amount of light allowed through the pupillary opening neural adaptation, involving the neurons in the successive stages of the visual chain in the retina itself and in the brain degree of adaptation fewfold,but occurs in a fraction of a second

Importance of Dark and Light Adaptation

Between the limits of maximal dark adaptation and maximal light adaptation, the eye can change its sensitivity to light as much as 500,000 to 1 million times, the sensitivity automatically adjusting to changes in illumination. Because registration of images by the retina requires detection of both dark and light spots in the image, it is essential that the sensitivity of the retina always be adjusted so that the receptors respond to the lighter areas but not to the darker areas. It

example of maladjustment of retinal adaptation Enter the sun from a movie theater, even the dark spots appear bright leaving little contrast. Enter darkness from light, the light spots are not light enough to register. the intensity of sunlight is about 10 billion times that of starlight, yet the eye can function both in bright sunlight after light adaptation and in starlight after dark adaptation.

Sensitivity and Acuity

Rods and cones synapse with bipolar cells Bipolar cells synapse with ganglion cells Ganglion cells synapse with neurone fibres At the fovea each cone synapses individually with a ganglion cell This gives good Acuity (resolution).

Many Rods synapse with one bipolar neurone RETINAL CONVERGENCE

Dim light results in small amount of neurotransmitter release Individually, this would be insufficient to over come the threshold of the bipolar cell, but the total amount of transmitter from several rods is sufficient. This gives less acuity but better sensitivity

Photochemistry of Vision

Rods and cones contain chemicals that decompose on exposure to light. This excites the nerve fibers leading from the eye. The membranes of the outer-segment of the rods contain rhodopsin or visual purple. Rhodopsin is a combination of a protein called scotopsin and a pigment, retinal. The retinal is in the cis configuration. Only the cis configuration can bind with scotopsin to form rhodopsin.

Light and Rhodopsin

When light is absorbed by rhodopsin it immediately begins to decompose. Decomposition is the result of photoactivation of electrons in the retinal portion of rhodopsin which leads to a change from the cis form of the retinal to the trans form of the molecule. Trans retinal has the same chemical structure but is a straight molecule rather than an angulated molecule. This configuration does not fit with the binding site on the scotopsin and the retinal begins to split away. In the process of splitting away a number of intermediary compounds are formed.

Mechanism for Light to Decrease Sodium Conductance

cGMP is responsible for keeping Na+ channel in the outer segment of the rods open. Light activated rhodopsin (metarhodopsin II) activates a Gprotein, transducin. Transducin activates cGMP phosphodiesterase which destroys cGMP. Rhodopsin kinase deactivates the activated rhodopsin (which began the cascade) and cGMP is regenerated reopening the Na+ channels.

Role of Vitamin A

Vitamin A is the precursor of all-transretinal, the pigment portion of rhodopsin. Lack of vitamin A causes a decrease in retinal. This results in a decreased production of rhodopsin and a lower sensitivity of the retina to light or night blindness.

The Rod Receptor Potential

Normally about -40 mV Normally the outer segment of the rod is very permeable to Na+ ions. In the dark an inward current (the dark current) carried by the Na+ ions flows into the outer segment of the rod. The current flows out of the cell, through the efflux of K+, ions in the inner segment of the rod.

The Dark Current

In the dark an inward current (the dark current) carried by the Na+ ions flows into the outer segment of the rod.

Rod Receptor Potential (Contd)

When rhodopsin decomposes it causes a hyperpolarization of the rod by decreasing Na+ permeability of the outer segment. The Na+ pump in the inner segment keeps pumping Na+ out of the cell causing the membrane potential to become more negative (hyperpolarization). The greater the amount of light the greater the electronegativity.

The Dark Current

When rhodopsin decomposes in response to light it causes a hyperpolarization of the rod by decreasing Na+ permeability of the outer segment.

Color Vision

Color vision is the result of activation of cones. 3 types of cones: blue cone green cone red cone The pigment portion of the photosensitive molecule is the same as in the rods, the protein portion is different for the pigment molecule in each of the cones. Makes each cone receptive to a particular wavelength of light

the three types of cones show peak absorbencies at light wavelengths of 445, 535, and 570 nanometers,

Each Cone is Receptive to a Particular Wavelength of Light

orange light with a wavelength of 580 nanometers

stimulates the red cones 99 (99 percent of the peak stimulation at optimum wavelength); stimulates the green cones 42 , blue cones not at all . , the ratios of stimulation are 99:42:0.

set of ratios-0:0:97-is interpreted by the nervous system as blue.

ratios of 83:83:0 are interpreted as yellow , and 31:67:36 as green

. Perception of White Light About equal stimulation of all the red, green, and blue cones gives one the sensation of seeing white.

Color Blindness

Lack of a particular type of cone Genetic disorder passed along on the X chromosome Occurs almost exclusively in males About 8% of women are color blindness carriers Most color blindness results from lack of the red or green cones Lack of a red cone, protanope. Lack of a green cone, deuteranope.

Neural Organization of the Retina

Direction of light

Signal Transmission in the Retina

Transmission of signals in the retina is by electrotonic conduction. Allows graded response proportional to light intensity. The only cells that have action potentials are ganglion cells. send signals all the way to the brain

Lateral Inhibition to Enhance Visual Contrast

Horizontal cells connect laterally between the rods and cones and the bipolar cells Output of horizontal cells is always inhibitory Prevents the lateral spread of light excitation on the retina Have an excitatory center and an inhibitory surround Essential for transmitting contrast borders in the visual image

Lateral inhibition, the function of horizontal cells

Function of Amacrine Cells

About 30 different types Some involved in the direct pathway from rods to bipolar to amacrine to ganglion cells Some amacrine cells respond strongly to the onset of the visual signal, some to the extinguishment of the signal Some respond to movement of the light signal across the retina Amacrine cells are a type of interneuron that Aid in the beginning of visual signal analysis.

Rods, Cones and Ganglion Cells

Each retina has 100 million rods and 3 million cones and 1.6 million ganglion cells. 60 rods and 2 cones for each ganglion cell At the central fovea there are no rods and the ratio of cones to ganglion cells is 1:1. May explain the high degree of visual acuity in the central retina

Three Types of Ganglion Cells

W cells (40%) receive most of their excitation from rod cells. sensitive to directional movement in the visual field X cells (55%) small receptive field, discrete retinal locations, may be responsible for the transmission of the visual image itself, always receives input from at least one cone, may be responsible for color transmission. Y cells (5%) large receptive field respond to instantaneous changes in the visual field.

Excitation of Ganglion Cells

Spontaneously active with continuous action potentials Visual signals are superimposed on this background Many excited by changes in light intensity Respond to contrast borders, this is the way the pattern of the scene is transmitted to the brain

Eye Movements are Controlled by 3 Separate Pairs of Muscles.

Superior and inferior obliques rotate the eyes Medial and lateral recti move eyes side to side Superior and inferior recti move eyes up and down
Figure 51-7; Guyton & Hall

Neural Pathways for Control of Eye Movement

Fixation movements of the eyes controlled by two neuronal mechanisms, voluntary and involuntary.
Voluntary fixation movements controlled by an area in the premotor cortex. Involuntary fixation mechanism causes eyes to lock on object of attention found with the voluntary fixation mechanism. Controlled by secondary visual areas of the occipital cortex. Results from negative feedback mechanism controlled at the level of the superior colliculus that prevents objects of attention from leaving the foveal portion of the retina.

Neural Pathways for Control of Eye Movement

Figure 51-8; Guyton & Hall

Saccadic Eye Movements

When the visual scene is moving (turning the head), the eyes fix on one highlight after another in the visual field jumping at a rate of 2 to 3 jumps/sec. These jumps are called saccades, and the movements are called opticokinetic movements. Saccades occur very rapidly (only 10% of the time is spent making saccades). Vision is suppressed during a saccadic movement.

The Autonomic Nerves to the Eyes

The eye is innervated by both parasympathetic and sympathetic neurons. Parasympathetic fibers arise in the EdingerWestphal nucleus, pass in the 3rd cranial nerve to the ciliary ganglion.
Postganglionic fibers excite the ciliary muscle and sphincter of the iris.

Sympathetic fibers originate in the intermediolateral horn cells of the superior cervical ganglion.

Postganglionic fibers spread along the corotid artery and eventually innervate the radial fibers of the iris.

Autonomic Pathways to the Eye

Figure 51-7; Guyton & Hall

Control of Accommodation (Focusing the Eyes)

results from contraction or relaxation of the ciliary muscle regulated by negative feedback mechanism that automatically adjust the focal power of the lens for highest degree of visual acuity within about 1 sec exact mechanism is not known

Control of Pupillary Diameter

miosis: decreasing of pupillary aperture due to stimulation of parasympathetic nerves that excite the pupillary sphincter muscle mydriasis: dilation of pupillary aperture due to stimulation of sympathetic nerves that excite the radial fibers of the iris

Pupillary Light Reflex

When the amount of light entering the eyes increases, the pupils constrict. Functions to help the eye adapt extremely rapidly to changing light conditions. Light excites fibers going to pretectal nuclei. From pretectal nuclei fibers pass to EdingerWestphal nucleus and back through parasympathetic nerves to constrict iris sphincter.

Visual field of left eye Temporal half Nasal half Nasal half

Visual field of right eye Temporal half

Left eye Temporal retina Nasal retina

Right eye Nasal retina

1 2

3 4 Optic 4
tract

3 5

1 2

Temporal retina

Midbrain

5
Lateral geniculate nucleus of the thalamus

Midbrain

Optic radiations

Optic radiations

Primary visual area of cerebral cortex (area 17) in occipital lobe Left eye and its pathways Right eye and its pathways