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Physiology of Vision: Angelie Rose C. Guzman, Rph. MD
Physiology of Vision: Angelie Rose C. Guzman, Rph. MD
Physiology of Vision: Angelie Rose C. Guzman, Rph. MD
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Lecture Outline
I. Structure of the Eye IV Adaptation
A. Light Adaptation
A. Outer Layer B. Dark Adaptation
1. Conjunctiva 1. Dark Adaptation Curve
2. Purkinje Shift
2. Cornea V. Phenomenon of Diffusion
3. Sclerae VI. Rods and Cones
VII. Color Vision
B. Middle Layer
A. Spectral Sensitivity under Dark- and Light-Adapted Conditions
1. Choroid B. Absorbance Spectra of Human Rods and Cones
2. Ciliary Body C. Color Sense
1. Classification of Color-Vision Defects
3. Iris VIII. Neuro-Ophthalmologic Optics
4. Additional Information A. Monocular Vision
B. Binocular Vision
C. Inner Layer 1. Panum’s Fusional Area
1. Retina C. Stereopsis (Depth Perception)
IX: Visual Pathway
II Photoreceptors
A. Parts of the photoreceptor
B. Types of Photoreceptors
C. Optic Nerve
III Visual Transduction/Phototransduction
A. Visual Transduction
B. Phototransduction
1. Dark Current
2. With Light
3. Absorption of Photons
C. Rhodopsin
D. Role of Calcium
E. Termination of Light-Activated State
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Structure of the eye
● Conjunctivae
Outer layer
● Cornea
(Fibrous coat)
● Sclerae
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A. OUTER/FIBROUS LAYER
1. CONJUNCTIVA
• Thin transparent mucous membrane that covers the
posterior surface of the lids (palpebral conjunctivae)
and the anterior surface of the sclerae (bulbar
conjunctivae)
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A. OUTER/FIBROUS LAYER
2. CORNEA
● Thin, transparent tissue inserted into the sclerae at the limbus
● Avascular, contains many pain fibers, and without pigmentation
○ Layers (ABCDE):
■ Anterior epithelium
■ Bowman's layer
■ Corneal stroma
● Thickest part of the cornea
● Composed of keratinocytes and collagen fibers
■ Descemet’s layer
■ Endothelium
● No mitotic activity
● Responsible for the relative deturgescence (relatively dehydrated state) of the cornea
● Absence of mitotic activity and dehydrated state is why the cornea is transparent 6
A. OUTER/FIBROUS LAYER
3. SCLERA
• Dense white fibrous outer protective coating consisting of collagen
• Continuous with the cornea (anteriorly) and the dural sheath of the optic nerve
(posteriorly)
• Attachment of the extraocular muscles
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Structure of the eye
● Conjunctivae
Outer layer
● Cornea
(Fibrous coat)
● Sclerae
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B. MIDDLE / VASCULAR LAYER (UVEAL TRACT)
1. CHOROID
• Highly vascular posterior segment of the uveal tract,
between the sclera and the retina
• Choriocapillaris: nourish the outer ⅓ of the retina
• Central retinal artery: nourishes the inner ⅔ of the
retina
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B. MIDDLE / VASCULAR LAYER (UVEAL TRACT)
2. Ciliary Body
• Triangular in cross section
• Extends forward from the anterior end of the
choroid to the root of the iris
• Two parts: ciliary muscle, ciliary processes
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Ciliary muscle
• Smooth muscle fibers with three arrangements:
■ Longitudinal, circular, radial
• When the muscles contract, the diameter of the ciliary
body reduces in size
Ciliary processes
• Attach the lens to the ciliary body via the zonular fibers
• Collectively, the zonular fibers form the suspensory
ligament of the lens
• Controls the shape of the lens
• Produces the aqueous humor
■ Circulates in the anterior chamber and maintains a
constant pressure inside the eye
■ Flow: from posterior chamber → pupil → anterior
chamber → trabecular meshwork → canal of
Schlemm → episcleral vessels → systemic circulation
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3. IRIS
• Anterior extension of the
ciliary body
• Separates the anterior and
the posterior chamber
• Flat surface with a centrally
situated round aperture
known as the pupil
• Within the stroma of the iris
are the sphincter (circular)
and dilator (radial) muscles
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Pupil Size Pathways
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A. ACCOMMODATION
• The change in the refractive power of the eye
to focus on near object
• Control is parasympathetic > sympathetic
• Process
1. Contraction of the ciliary muscle
2. Relaxation of the lens capsule
3. Increased anteroposterior diameter of
the lens
4. Miosis (pupillary constriction
5. Convergence
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B. Refraction
● Refraction: the bending of light rays at an angulated
interface
● Optical components of the eye collect light and focus the
image on the retina (light → iris → pupil → lens → retina)
Index of Refraction
● Essentially a measure of the speed of light within it
○ Light travels faster through air than through the cornea
● Determinants of how much a light ray is refracted:
1. Difference in the refractive indices of the two
media (ex. between the refractive indices of air and
of the cornea)
2. Angle between the incident light and the interface
between the two media
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• Emmetropia
● Normal vision
● Parallel light rays from
distant objects are in
sharp focus on the
retina when the ciliary
muscle is completely
relaxed
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Errors of Refraction
• Myopia
○ Also termed as nearsightedness
○ The images of distant objects are
focused in front of the retina
○ Usually due to too long an eyeball
○ Corrected by using biconcave lenses
• Hyperopia
○ Also termed as farsightedness
○ The images are focused at the back of
the retina
○ Due to either a short eyeball or a
weak lens system
○ Corrected by using convex lenses
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Errors of Refraction
• Astigmatism
○ Causes the visual image in one plane to focus at
a different distance from that of the plane at
right angles.
• A point source of light cannot be brought to
precisely focus on the retina
○ Usually results from too great a curvature of the
cornea in one of its planes
• Uneven curvature of the refractive surfaces of the
eye
○ Corrected by using spherical or cylindrical
lenses that will correct for both axis and strength
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Presbyopia
• The lens continues to enlarge throughout life
■ As we age, the lens grows larger and thicker and becomes less elastic partly
Errors of
because of progressive denaturation of lens proteins
↑ Age = ↑ Size & Thickness of Lens = ↓ Elasticity
Refraction • Example: the elderly having a hard time looking at their cellphones because they cannot
clearly see near objects
• Corrected by using bifocal lenses
■ Upper segment: used to see far objects
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■ Lower segment: used to see near objects
• Vitreous Humour
● Clear, avascular
gelatinous body that fills the
space between the lens and
retina
● 99% water, and 1%
collagen and hyaluronic acid
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Structure of the eye
● Conjunctivae
Outer layer
● Cornea
(Fibrous coat)
● Sclerae
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C. INNER/NEURAL LAYER
1. RETINA
● Embryogically and histologically a part of the CNS
● Transduces light into neural signals and does
complex processing of visual information before
passing it in the different regions of the brain
● Thin sheet of tissue that lines the back of the eye
● Contains light sensitive cells known as the
photoreceptors
○ Capture photons, converts light energy to
chemical free energy as signals to the visual
neurons in the retina
● Contains the macula, wherein at its center, there is
a pit-like depression known as the fovea 26
• Macula
○ Where visual acuity is sharpest
○ Most essential part of the retina for human vision
○ Highest concentration of cones
• Fovea
○ Pit-like depression in the middle of the macula
○ Where visual targets are most focused
○ Foveal vision is purely cone-mediated
○ Only cones are present
■ Light passes unimpeded to the cones
● Blood vessels, ganglion cells, the inner
nuclearlayers of cells and the plexiform layers are
all displaced to one side rather than resting directly
on top of the cones
■ Each cone photoreceptor is synapsed to a single
bipolar cell, which in turn synapse to a single ganglion
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cell
• Peripheral Vision
○ Visual angle >10° away from the fovea
○ Vision is mediated by both rods and cones
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Layers of the Retina
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• Interneurons: Synaptic connections within the retina
■ Bipolar cells
● Directly connect photoreceptors and ganglion
cells radially
■ Horizontal cells
● Mediate interactions within the wide area of
the retina
● Synapse within the outer layer of the retina
● Interconnect photoreceptors and bipolar cells
to themselves and to each other
■ Amacrine cells
● Synapse within the inner layer of the retina
● Interconnect both bipolar and ganglion cells 30
● Retinal Pigment Epithelium
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Summary
Outer Conjunctiva Thin, transparent mucous membrane
Iris Separate ant and post chamber; with central round aperture – pupil
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Pleaseeeeeee……
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II. PHOTORECEPTORS
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B. TYPES OF PHOTORECEPTORS
● Rods
○ Extremely sensitive and responds to very low levels of illumination
○ Outnumber the cones by at least 16:1
○ Responsible for scotopic vision (operates best under reduced lighting)
○ Outer segment: has about 1000 stacks of disk membranes
■ Flattened, membrane bound organelles that have pinched
off from the outer membrane
■ Float freely in the outer segment after being disconnected
from the plasma membrane when formed at the base
■ The disk membrane contain photopigment called the
rhodopsin (retinal and scotopsin)
■ Human retina has approximately 120 million rods
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B. TYPES OF PHOTORECEPTORS
● Cones
○ Respond to light brighter than twilight
○ Less sensitive than rods
○ Responsible for photopic vision
○ High visual acuity and color vision
○ Outer segment: has stacked disk membranes but they are infolded and
remain continuous with the plasma membrane unlike the rods
○ The disk membrane contain photochemicals (retinal and photopsins)
○ Cone density falls to very low levels outside the fovea and rod density rises
○ Human retina has about 6 million cones
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Differences between Rods and Cones
Rods Cones
High sensitivity; specialized for night vision Lower sensitivity; specialized for day vision
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III. PHOTOTRANSDUCTION
1. Dark Current
2. With light
3. Absorption of photons
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1. DARK CURRENT
● This is carried mainly by inwardly directed Na+ in the outer segment and by outwardly
directed K+ ions in the inner segment
● Each photoreceptor produces an ionic current that flows steadily into the outer
segment and out of the inner segment
● Na+ flows inwardly through a nonselective cation channel of the outer segment which
light indirectly regulates K+ flows through a potassium channel in the inner segment
which light does not regulate
● Na-K pumps: primarily located within the inner segment, removes Na+ and import K+
● Na-Ca exchanger: removes Ca2+ from the outer segment
● Na+ carries ~90% of the dark current in the outer segment and Ca2+ carries ~10%
● There is release of the transmitter glutamate, when presynaptic terminal is depolarized
● In the dark, cGMP levels are high hence more cGMP-gated cation channels are open
● When cGMP-gated cation channels are open, Na+ ions influx causing the depolarization
of the photoreceptor
● As the photoreceptor is depolarized, neurotransmitter release is high and this is called
the dark current
● In the absence of light, the dark current, which depolarizes the cell, leads to constant
transmitter release.
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● In the dark, there is continuous supply of
cGMP synthesized from GTP with the help of
guanylyl cyclase
● High cytoplasmic cGMP level causes cGMP-
gated non-selective cation channels to open
leading to Na+ influx and depolarization
● This is known as the dark current
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2. WITH LIGHT
● In the presence of light, Na+ can no longer enter because
cGMP levels are low
● The cGMP-gated channels are closed, while K+ channels
in the inner segment remains open
● K+ continuously leak out of the inner segment causing
the photoreceptor cell to hyperpolarize
● When the photoreceptor is hyperpolarized, the
neurotransmitter release decreases
● At low light intensities, the size of the receptor potential
rises linearly with light intensity
● At the higher intensities, the response saturates
● A flash of light causes a decrease in transmitter secretion,
meaning that the photoreceptor is most active in the dark
● In the presence of light, the photoreceptor cell thus
hyperpolarizes, and transmitter release decreases.
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• In the presence of light, transducin is activated by
metarhodopsin II, the alpha subunit of transducin
exchanges a bound GDP for a GTP which then
stimulates phosphodiesterase
• Phosphodiesterase hydrolyzes cGMP to 5’ GMP leading
to a decreased concentration of cGMP
• The decreased cGMP level closes the cGMP-gated non-
selective cation channel leading to the
hyperpolarization of the photoreceptor.
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3. ABSORPTION OF PHOTONS
● Leads to closure of the nonselective cation channels in the outer segment
● Because the K+ channels of inner segment remains open, this outward current cause
the cell to hyperpolarize
● Number of the cation channels that close depends on the number of photons that
are absorbed
○ Sensitivity of the rods is 1 to approximately 1000 photons
○ Cones are less sensitive, but they are faster than rods
○ Cone responses do not saturate even at the brightest levels of
natural light
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C. RHODOPSIN
● G protein coupled light receptor
● A single photon can stop the flow of a million Na+
ions across themembrane of a rod cell. The photon is
absorbed by rhodopsin, the light receptor molecule.
● Its molecule is made up of retinal and opsin
● Opsin is a single polypeptide with 7 membrane
spanning segments
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● Electrons of the retinal are photoactivated and 11 cis-
retinal is isomerized to all trans-retinal
● The straight configuration of the all trans-retinal renders
the retinal molecule ill-fitting with the reactive sites of
opsin hence they pull apart from each other and it is
known as bleaching
● As retinal detaches from its opsin molecule,
metarhodopsin II is formed
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● As 11 cis-retinal is isomerized to all trans-retinal it
produces metarhodopsin II
● Metarhodopsin II stimulates transducin that
activates phosphodiesterase which hydrolyzes cGMP to
5’-GMP
● Low levels of cGMP, closes the cGMP cation channels
thus decreasing Na+ influx
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SUMMARY:
PHOTOTRANSDUCTION (in the dark/dark current)
● There is continuous supply of cGMP synthesized from GTP by guanylyl cyclase
● Increased cytoplasmic cGMP level opens cGMP-gated non-selective cation channels
● This leads to Na+ influx and depolarization of the photoreceptor
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SUMMARY:
PHOTOTRANSDUCTION (in the dark/dark current)
● ↑ Guanylyl cyclase activity ; ↑ cGMP concentration
● opens cGMP-gated non-selective cation channels
● Na+ influx = depolarization
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D. ROLE OF CALCIUM
● cGMP-gated non-selective cation channel
○ Allows not only Na+ ions to enter but also allows calcium
to enter the outer segment of the photoreceptor, hence the
term “non-selective”
○ Calcium that enters sends a negative feedback mechanism,
thus regulating the level of cGMP
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1. WITHOUT LIGHT
● Absence of light causes continuous stimulation of
guanylyl cyclase and inhibition of
phosphodiesterase
● This leads to an increased level of cGMP
● Increased cGMP opens cGMP-gated cation
channels, allowing Na+ and Ca2+ to enter
● Calcium that enters inhibits guanylyl cyclase,
decreasing cGMP production and also slightly
stimulates phosphodiesterase to decrease cGMP
concentration.
● This will close the non-selective cation channel.
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2. WITH LIGHT
● Presence of light causes inhibition of guanylyl cyclase and
activation of transducin, leading to stimulation of
phosphodiesterase
● This leads to a decreased level of cGMP
● Decreased cGMP closes cGMP-gated cation channels
● Remaining calcium inside the cytosol will exit via the Na-Ca
Exchanger
● The decreased calcium level inside will stop the inhibition of
guanylyl cyclase, leading to more production of cGMP
● The decreased calcium level will also inhibit phosphodiesterase,
leading to more production of cGMP
● This leads to priming of the cGMP-gated cation channels to
reopen
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SUMMARY:
ROLE OF CALCIUM (without light) ROLE OF CALCIUM (with light)
● ↑ Guanylyl cyclase activity ; ↑ cGMP concentration ● ↓ Guanylyl cyclase activity ; ↓ cGMP concentration
● ↓ Phosphodiesterase activity ; ↑ cGMP concentration ● ↑ Transducin activation ; ↑ Phosphodiesterase activity
● ↑ cGMP concentration ; ↑ Opening of cGMP-gated ● ↑ Phosphodiesterase activity ; ↓ cGMP concentration
cation channels
● ↓ cGMP concentration ; ↑ Closing of cGMP-gated cation
● ↑ Opening of cGMP-gated cation channels ; ↑ Na and channels
Ca influx
This will lead to:
● ↑ Cytosolic calcium ; ↓ guanylyl cyclase activity
● Exit of calcium via Na-Ca exchanger
● ↑ Cytosolic calcium ; ↑ Phosphodiesterase (slightly)
● ↓ Cytosolic calcium ; ↓ inhibition of guanylyl cyclase (will
stop inhibition)
Therefore, ● ↓ Cytosolic calcium ; ↓ phosphodiesterase
● ↑ Cytosolic calcium ; ↓ cGMP concentration
(via inhibition of guanylyl cyclase and stimulation of Therefore,
phosphodiesterase)
● ↓ Cytosolic calcium ; ↑ cGMP concentration
● ↑ cGMP concentration ; ↑ Priming of cGMP-gated cation
channels to reopen 55
E. TERMINATION OF LIGHT-ACTIVATED STATE
● May be from
○ Closure of cGMP-gated channels, leading to
decreased calcium level
○ Rhodopsin kinase
■ Phosphorylates light-activated rhodopsin
■ Acts on metarhodopsin II (which is
responsible for activating transducin)
○ Recognition by cytoplasmic protein known as
Arrestin
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IV. ADAPTATION
A. LIGHT ADAPTATION
● When we move from total darkness to full light
● Occurs rapidly over a few minutes
● Cones subserve daylight or photopic vision, color vision, contour
resolution, or visual acuity
B. DARK ADAPTATION
● When we move from full bright light to a dark environment
● Requires longer period of time (>30 mins)
● Rods subserve mainly night or scotopic vision and have no color perceptive
ability
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Initially, upon entering a dark movie theater, you will have difficulty seeing
the surroundings, but you will see your surroundings better after a few
minutes.
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1. DARK ADAPTATION CURVE
● When coming from a bright environment to a dark environment, the retinal
sensitivity is initially low
● Cones
○ Adapt more rapidly than the rods (early/left portion of the curve)
○ Cease to adapt after a few minutes with decreased retinal sensitivity
● Rods
○ Slow-adapting
○ Continue to adapt for a longer period with increased retinal sensitivity
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2. PURKINJE SHIFT
● Occurs at the transition between primary use of photopic to scotopic systems, and
vice versa
● As light intensity decreases, the rods take over and before the color disappears,
there is full sensitivity of the rods (we are able to adapt to the dark environment)
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V. PHENOMENON OF DIFFUSION
● The phenomenon of diffusion occurs when
cones at the center of a light beam receives the
highest energy, while the peripheral cones
receive less energy
● The beam of light is not focused at a point but
as heaps of energy
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VII. COLOR VISION
● Color sensitive pigments are made up of retinal and photopsins
● Sensitivity to light wavelength is dependent on retinal adaptation
○ Scotopic or dark adapted
■ sensitivity curve shifts towards the shorter wavelengths
○ Photopic or light adapted
■ sensitivity curve shifts towards the longer wavelengths
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A. SPECTRAL SENSITIVITY UNDER DARK- &
LIGHT-ADAPTED CONDITIONS
● If we plot the visible spectrum according to the
wavelength of light and luminous efficiency
(minimum intensity of the wavelength to produce a
response), we would get a spectral sensitivity curve.
● The absorption spectrum of rods rhodopsin peaks
at 500 nm
● The absorption spectrum of cones pigments peaks
at 560 nm
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B. ABSORBANCE SPECTRA OF HUMAN
RODS AND CONES
● Short (S) wavelength that peaks at 420 nm
represents the BLUE cones
● Medium (M) wavelength that peaks at 530 nm
represents the GREEN cones
● Long (L) wavelength that peaks at 560 nm
represents the RED cones
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C. COLOR SENSE
● This is the function of the cone photoreceptors in addition to the form sense.
● Color defective vision occurs in about 10% of males and almost 1% of females as
sex-linked recessive hereditary condition.
● Truly color-blind individuals are rare.
○ These are individuals with total absence of one type of photo pigment or
color sensitive cone.
● Anomaly – relative deficiency of one color.
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1. CLASSIFICATION OF COLOR-VISION DEFECTS
● Hereditary
● Monochromats – has only one color pigment and completely lacks the ability to distinguish wavelengths in the spectrum
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VIII. NEURO-OPHTHALMOLOGIC OPTICS
A. MONOCULAR VISION
● In monocular vision, all visual impressions from the left side cross over to the right cerebral cortex and vice versa.
B. BINOCULAR VISION
● The ability to combine the visual information from two eyes viewing the same scene into a single image.
○ The visual impressions from the left side of the visual field still cross over to the right visual cortex and vice
versa.
○ The only difference from monocular vision is that both retinas send the same visual impressions in the visual
cortex. This means that the visual impression of the left half of the field falling on the left retina and the same
impression falling on the right retina both go to the right visual cortex.
○ The impression falling on the left retina has to cross over to go to the right cortex while that from the right
retina has no need to cross.
● This is because of Sensory Fusion
• ○ The process whereby disparities between the two images at corresponding retinal points are overcome to
allow a single image to be perceived 70
1. PANUM’S FUSIONAL AREA
● The fusible point is called the Panum’s
Fusional Area
● The horopter is a point in space which
projects on corresponding points in the two
retinas that are on anatomically identical
points
● The Panum’s Area is the region in front and
back of the horopter in which single vision is
present
○ This is also known as the Panum’s area of
single binocular vision.
● This means that if an image falls within the
Panum’s Fusional Area, it would be perceived
as a single image
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C. STEREOPSIS (DEPTH PERCEPTION)
● Each point in the retina has a corresponding point in the retina of the other eye
and the visual impression from both points end up in the same point in the visual
cortex.
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Visual Pathway
● Fibers of the optic tracts synapse in the
dorsal lateral geniculate nucleus of the
thalamus
● Fibers pass via the optic radiation to go to
the primary visual cortex
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End..
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