Physiology of Vision

Zoology

Lesson: Physiology of Vision

Lesson Developer: Dr. Mahtab Zarin

College/Dept: Zoology, University of Delhi

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 Physiology of Vision

Table of Contents
 Introduction
 Image formation mechanisms
 Refraction of light rays
 Accommodation
 Change in pupil size
 Types of vision
 Errors of refraction
 Photochemistry of vision
 Rhodopsin-retinal visual cycle and excitation of rods
 Processing and transmission of visual impulse
 Visual perception
 Summary
 Exercises
 Glossary
 References

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Learning objectives
 To describe the structure of eyes as a photoreception organ
 Pathways of Visual information from eye to brain
 To understand the mechanisms of image formations
 Role of the photoreceptors in image processing
 Refraction of the light rays on cornea
 To understand the visual perception of the object

Introduction

Vision is the special sense of sight that is based on the transduction of light stimuli received
through the eyes. Each eye with layer of receptors, lens system, and nerves act as sensory
receptor for vision.

Receptors of the eyes are able to detect a small portion of the vast spectrum of the
electromagnetic radiation that we call visible light (Fig. 5). The wavelengths capable of
stimulating the receptors of the eye-visible spectrum- are between about 400 and 700nm.
Different wavelengths of light within band are perceived as different colors.

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Fig. 1. Electromagnetic spectrum

Visible light ranges in wavelength from 400 to 700nm.
Source: http://en.wikipedia.org/wiki/File:EM_spectrum.svg
Image Credit: This file is licensed under the Creative Commons Attribution ShareAlike 3.0 License.

Value addition: Did you Know

Color of Object depends on the wavelength reflected
An object can absorb certain wavelengths of visible light and reflect others; the
object will appear the color of wavelength that is reflected. For example,
 A green apple appears green because it reflects mostly green light and absorbs
most other wavelengths of visible light.
 An object appears white because it reflects all wavelength of visible light.
 An object appears black because it absorbs all wavelength of visible light.
Source: Principles of Anatomy & Physiology- Tortora, G.J. & Derrickson, B.

Image formation mechanisms

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To create clear vision, light reflected from objects within the visual field is focused on to the
retina of each eye. The followings are the steps implicated in achieving a clear image:
1. the Refraction or bending of light through lens and cornea in eye;
2. Accommodation, the adjustment in shape of the lens;
3. Changing the size of the pupils, Constriction or narrowing

Refraction of light rays

Refraction is of light rays is bending of the rays at an angulated interface. It is a
phenomenon that often occurs when light rays travel from a medium with a given refractive
index to a medium with another at an oblique angle. At the boundary between the media,
the wave's phase velocity is altered, usually causing a change in direction. Its wavelength
increases or decreases but its frequency remains constant. The degree of refraction
increases as a function of
i) ratio of the two refractive indices of the two transparent media
ii) degree of angulation between the interface and the entering wave front.
Refractive index can be defined as the ratio of velocity of light in air to the velocity in a
substance.
Human eye is optically equivalent to a camera which has a lens system, a variable aperture
in form of pupil and a retina which corresponds to photographic film.
Lens system of the eye has four refractive interfaces between:
a) air and anterior surface of cornea
b) posterior of camera and aqueous humor
c) aqueous humor and anterior surface of lens
d) posterior surface of lens and vitreous humor.
The refractive index of air is 1.00, cornea-1.38, aqueous humor-1.33, lens-1.40 and
vitreous humor-1.34.

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Fig.2. Refractive indices of various parts of eye

Source:
http://cnx.org/resources/e7896dbfd44b4103c503e3727bd94a25/Figure_27_01_01.j
pg
ILLL in house
If we consider the Lens system of the eye as one unit and add up all the refractive indices,
the system would become simplified and the eye would be called as reduced eye. The
reduced eye is an idealized model of the optics of the human eye. Introduced by Franciscus
Donders, the reduced eye model replaces the several refracting bodies of the eye (the
cornea, lens, aqueous humor, and vitreous humor) are replaced by an ideal air/water
interface surface that is located 20 mm from a model retina. The single refractive surface
has its central point 17mm in front of retina and a total refractive power of 59 diopters
when the lens is accommodated for distant vision. About two third of the refractive power of
the eye is provided by the cornea because the refractive index of cornea is markedly
different from the air whereas the refractive indices of rest of lens system are not greatly
different.
The total refractive power of the internal lens of the eye is only 20 diopters, about one third
the total refractive power of eye

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Value addition: Did you Know

Diopters: a measuring unit of refractive power

The greater the curvature of a lens , the greater its refractive power. The refractive
power of a lens is conveniently measured in diopters, the number of diopters being
the reciprocal of the principal focal distance in meters. For example , a lens with a
principal focal distance of 0.25m has a refractive power of 1/0.25, or 4 diopters. The
human eye has a refractive power of approximately 66.7 diopters at rest.

Source: Review of Medical Physiology. Ganong, William F.

Accommodation
The lens of human eye is biconvex i.e it is convex on both its anterior and posterior
surfaces. Its ability to refract light increases as its curvature becomes greater. The change
in the shape of eye lens let the focusing power of eyes adjust for far and close vision. It
differs with the extent of light which is to be refracted (bent). In order to produce a sharp
image on the retina, light rays from close objects must be refract more than those from far
placed objects. This phenomenon is known as accommodation.
Accommodation for close vision
In order to focus on near objects i.e. within about 6 meters, accommodation is required and
the eye must make the following adjustments
 Constrictions of the pupils
 Convergence
 Changing the power of lens
Constrictions of the pupil: This assists accommodation by reducing the width of the beam
of light entering the eye so that that it passes through the central curved part of the lens.
Convergence (movement of eyeballs): Extrinsic muscles help to move the eyes. To
achieve a sharp image, the eyeball has to be rotated so that they converge on the object
viewed. The closer an object is to the eyes, the more eye rotation is needed to accomplish
convergence.
Changing the power of lens: Changes in the thickness of the lens are made to focus light
on the retina. The amount of adjustment depends on the distance of the object from the
eyes. When you view a close object, the ciliary muscle contracts which pulls the ciliary body
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and choroid forward toward the lens. As a result the lens becomes more convex, which
increases its focusing power.
Accommodation for distant vision
When objects more than 6 metres away, the ciliary muscle of the ciliary body is relaxed and
lens is flatter because it is stretched in all directions (Fig.3).

Fig. 3.

Accommodation by human eye

Source: ILLL in house

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Fig. 4. Accommodation for near and distant object

Source: ILLL in house
Mechanism of accommodation and its control
In young person, the lens is composed of a strong elastic capsule filled with viscous,
proteinaceous but transparent fluid. When in relaxed state with no tension at all, lens
assumes a spherical shape due to elastic retraction of lens capsule. Suspensory ligaments
attach radially around the lens, pulls the lens towards the edges on outer surface of the
eyeball. The tension on these ligaments keeps the lens relatively flat under normal
conditions of eye.
The ciliary muscles located laterally to the lens ligaments have two separate set of smooth
muscle fibers. The contraction of either of these muscle fibers releases the ligaments
tension on the lens and the lens assumes a more spherical shape.
The ciliary muscle movement is controlled by parasympathetic nerve signals transmitted to
the eye through the third cranial nerve from third nucleus of brain stem. Stimulation of
parasympathetic nerves contracts both sets of ciliary muscles which relax the lens ligament
and hence lens becomes thicker and refractive power increased.
Presbyopia

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With growing age, especially after 40, the lens grows larger and thicker and becomes less
elastic because of partial denaturation of lens proteins. Hence the ability to change lens
shape decreases and so is the power of accommodation. The power of accommodation
decreases from about 14 diopters (in childhood) to 2 diopters at age of 45-50. It becomes 0
diopters after 70 when the lens becomes totally nonaccomodating. This condition is called
Presbyopia. Each eye remains focused permanently at an almost constant distance and can
no longer accommodate for both near and distant vision. Bifocal glasses are used to treat
this condition.

Change in the pupil size

The major function of the iris is to increase or decrease the amount of light that enters eye
during darkness or brightness. The amount of light that enters the eye though the pupil is
proportional to the area of the pupil (diameter 2). The pupil can become as small as 1.5mm
or as large as 8 mm in diameter.
Size of pupil affects the accommodation via allowing the required amount of light to enter
inside the eyeball. In dim light, the pupil size increases (means dilates) to let the maximum
amount of light to arrive at the light receptive retina. In bright condition, the pupil size
decreases (that means it constricts).
The range of image distances over which the image of an improperly focused object is
acceptably sharp is called the depth of focus. Depth of focus of lens system increases with
decreasing pupillary diameter. This is because with a very small aperture, almost all the
rays pass through the center of the lens and the central most rays are always in focus. In
other words, to have a sharp image, the pupillary diameter must be adjusted as per
distance of the object from eye or the image might become blurred.

Types of vision
With both the eyes open, the outer region of our total visual field is perceived by only one
eye known as zone of monocular vision. In the central portion, the fields from the two
eyes overlap known as the zone of binocular vision (Fig 5).
Parallel processing of information continues all the way to and within the cerebral cortex to
the highest stages of visual neural networks. Cells in this pathway respond to electrical
signals that are generated initially by the photoreceptors response to the light. Optic nerve
fibres projects to several structures in the brain, the largest number passing to the
thalamus) specifically toward the lateral geniculate nucleus of the thalamus, where the
information such as colour, intensity, shape and movement etc. from the different ganglion
cell types is kept distinct.
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Fig.5. Types of Vision

Source: http://caleblookeye.wikispaces.com/Stereoscopic+and+Monocular+Vision CC

Table 1. Difference between Monocular and Binocular vision

S.No. Binocular/ Stereoscopic Monocular Vision

vision
1. Gives a three-dimensional Monocular vision provides a limited
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image produced by two-dimensional view.

the fusion of two different views of
an object on each retina
2. In an animal with stereoscopic The visual fields of animals with
vision (e.g. humans), due to the monocular vision have little or no
overlapping of visual fields of both overlapping.
eyes , two eyes can focus on the
same object on retina. Though the
two eyes simultaneously produce
two images with little difference,
the brain resolves this into a three
dimensional impression which
therefore gives rise to stereoscopic
vision.
3. It gives a better judgement of the It helps to detect enemies from a wide
distance of the object range of directions.
4. Especially important Prey such as fishes, rabbits and
to predatory animals, such as grasshoppers usually have monocular
eagles, hawks, owls and cats, vision.
as good judgement of distance is
necessary to capture prey.
5. Animals with stereoscopic vision Animals with monocular vision have
have both eyes at the front of the eyes at the side of the head (laterally
head. However, the field of placed eyes). The field of vision of
vision of animals with stereoscopic animals with monocular vision
vision is narrow. is wide, but has restricted stereoscopic
vision.

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6.

Errors of refraction

Emmetropia (normal vision): An eye is said to be emmetropic if it can form a sharp

image of a distant object (at more than 6 meters) with all its ciliary muscles relaxed.
However to view the nearby objects, the eye must contract its ciliary muscle and thereby
provide appropriate degrees of accommodation.

Myopia (near-sightedness): It is a type of refractive error where image is formed in

front of retina for the light rays coming from distant object when all the ciliary muscles of
eyes are relaxed. This is the inability to see faraway objects clearly but have clear
nearsightedness. This can either be due to excessive curvature of refractive lens
(refractive myopia) or when the eyeball is longer in reference to the focusing power of the
lens (also known as axial myopia). Myopia is corrected by using a concave lens that
results in the deviation (divergence) of light rays prior to reaching the cornea.

Hyperopia (far-sightedness): It is a type of refractive error where image is formed

beyond retina for light rays coming from nearby objects when all the ciliary muscles of eyes
are relaxed. This is the inability to see near objects clearly but have clear farsightedness.
This can be either due to the weak focusing power as the length of the eyeball is short (also
known as axial hyperopia). This may also result from a cornea or crystalline lens with not
enough curvature (refractive hyperopia). It is corrected by convex lenses which cause
light rays to congregate (converge) before striking the cornea.

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Astigmatism: This is the defects of vision which occurs when the surface of cornea or
lens is not smoothly spherical. Due to this condition, some portions of image are out of the
focus, thereby the vision become blurred or altered. Cylindrical lenses can be used to
correct this condition.

Fig. 6. Refraction abnormalities and their correction

Source: ILLL in house

Photochemistry of vision
Once the eye is accommodated to see an object, all the transparent parts contribute to
converge the rays of light to form an image on the retina. An inverted image is formed on

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retina and the photoreceptor cells of retina finally perceive the image to convey it to the
CNS. The major events that follow are:
1. Rhodopsin-retinal visual cycle and excitation of rods
2. Processing and conduction of visual sensation, performed through the image
processing cells of retina and visual pathway, and
3. Perception of vision involves a role of visual cortex and associated regions of cerebral
cortex.

Rhodopsin-retinal visual cycle and excitation of rods

Visual cycle
 When light energy is absorbed by rhodopsin, the rhodopsin begins to decompose
within a fraction of seconds.
 Photoactivation of electron in the retinal component leads to instantaneous
conversion of cis form of retinal (angulated molecule) to all-trans form (straight
form).
 Because of geometric isomerization of retinal, the protein component scotopsin
cannot fit into the reactive site anymore. Hence it pulls away from the scotopsin.
 The immediate product is bathorhodopsin (partially split combination of all -trans
retinal and scotopsin).
 Being highly unstable, bathorhodopsin decays into lumirhodopsin.
 Subsequently it decays to metarhodopsin I followed by metarhodopsin II and
finally much more slowly into completely split products scotopsin and all-trans
retinal.
 So this portion of visual cycle (separation of opsin and retinal) is termed as
photodecomposition and the rhodopsin or photopigment is known to be bleached
through the action of lightor rhodopsin bleaching.
 Metarhodopsin II also called as activated rhodopsin excites the electrical changes
in rods and the rods then transmit the image to CNS through optic nerve. This
process is called Phototransduction.
 Reformation of rhodopsin: rhodopsin is reformed by conversion of all-trans retinal
to 11-cis retinal in presence of enzyme retinal isomerase. Once formed 11-cis retinal
automatically combines with scotopsin to form rhodopsin which remains stable until
it is decomposed by absorption of photon. the bleaching of the rhodopsin takes place

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in the presence of light, whereas the regeneration event occurs in the absence of
light
 This completes the visual cycle. Under equilibrium state, the rate at which the
photopigments are being bleached must be equal to the rate at which they are
regenerated.

A.

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B.
Fig.7 a. Diagrammatic representation of Rhodopsin-retinal visual
cycle
7 b. Rhodopsin-retinal visual cycle through flowchart
Source: ILLL in house
Value addition: Did you Know??

Heading text: Why Vitamin A deficiency causes Night blindness?

Body text: The severe deficiency of Vitamin A in a person causes night Blindness.
Vitamin A is normally present in both cytoplasm of rod and pigment layer of retina.
Therefore Vitamin A is normally always available to form new retinal in retina.
Deficiency of Vitamin A leads to severe depression of system for formation of retinal
and rhodopsin. Due to lack of rhodopsin and retinal, the person is unable to see in

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dim light as the light is too low to excite reaction.

Source: Textbook of Medical Physiology. Guyton and Hall

Excitation of rods
When rhodopsin is activated by light, it causes excitation of rod. Excitation of rod causes
increased negativity of intrarod membrane potential, which is a state of hyperpolarization
(inside the negativity is more as compared to normal condition). Rhodopsin increases the
negativity by decreasing the rod membrane conductance for sodium ions in the outer
segment of the rod leading to hyperpolarization of entire rod membrane. Photoreceptor cells
are unusual cells in that they depolarize in response to absence of stimuli or scotopic
conditions (darkness). In photopic conditions (light), photoreceptors hyperpolarize to a
potential of -60mV. It is this 'switching off' that activates the next cell and sends an
excitatory signal down the neural pathway.

Value addition: Interesting to know

Heading text: Visual Transduction

Body text: The entire phenomenon of transforming light rays (refracted) into nerve
impulse is called as phototransduction. Visual phototransduction is a process by
which light is converted into electrical signals in the rod cells, cone cells and
photosensitive ganglion cells of the retina of the eye.
In the light
Light transduces the visual pigment via the following enzyme cascade: photons –
rhodopsin – activated rhodopsin (metarhodopsin II) –opsin activates the
regulatory protein a GTP binding protein (transducin) – transducin dissociates from
its bound GDP, and bind GTP, then the alpha subunit of transducin dissociates from
the beta and gamma subunits, with the GTP still bound to the alpha subunit–alpha
subunit-GTP complex activates cGMP-phosphodiesterase (an enzyme hydrolyzing
cGMP) –breaks down cGMP to 5'-GMP- lowers the concentration of cGMP and
therefore the sodium channels close –Closure of the sodium channels causes
hyperpolarization of the cell due to the ongoing potassium current–
Hyperpolarization of the cell causes voltage-gated calcium channels to close–
calcium level in the photoreceptor cell drops–the amount of the neurotransmitter
glutamate that is released by the cell also drops ( This is because calcium is required
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for the glutamate-containing vesicles to fuse with cell membrane and release their
contents).

Fig. 8a. Visual transduction in light

In the dark, a steady current flows into the open channels, carried mainly by Na
ions, constituting a ―dark current‖ {composed mainly of the influx of the Na+
component (80%) however, a Ca2+ component (15%) and a Mg2+ component
(5%)} that partially depolarizes the photoreceptor cell. Dark current keeps the cell
depolarized at about -40 mV. Thus, the depolarized photoreceptor releases
neurotransmitter (the amino acid glutamate) from its synaptic terminals upon
second-order neurons in the dark. On light stimulation the rhodopsin molecules are
isomerized to the active form, the above cascade ensues, leading to closure of the
cation channels of the photoreceptor membrane, stopping the dark current and
causing the photoreceptor cell membrane to hyperpolarize and cease
neurotransmitter release to second-order neurons.

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Fig. 8b. Visual transduction in dark

Deactivation of the phototransduction cascade
GTPase Accelerating Protein (GAP) interacts with the alpha subunit of transducin,
and causes it to hydrolyse its bound GTP to GDP, and thus halts the action of
phosphodiesterase, stopping the transformation of cGMP to GMP.
Click here to see an animation of phototransduction (Quicktime movie):
http://webvision.med.utah.edu/movies/trasduc.mov
Source:
 Textbook of Medical Physiology. Guyton and Hall
 http://en.wikipedia.org/wiki/Visual_phototransduction
 http://webvision.med.utah.edu/book/part-ii-anatomy-and-physiology-of-the-
retina/photoreceptors/
Image credit: ILLL in house

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Value addition: Video

Vision - Light and Neuronal Activity

<iframe width="420" height="315" src="//www.youtube.com/embed/AuLR0kzfwBU"

frameborder="0" allowfullscreen></iframe>

Source: You tube

Photochemistry of color vision

Different cones are sensitive to different colors of light. The absorption characteristics of the
pigments in the three types of cones blue, green and red show peak absorbencies at light
wavelengths of 445, 535 and 570 nm, respectively which helps the retina to differentiate
the colors.
The nervous system interprets the color on the basis of ratio of stimulation by
monochromatic light with a particular wavelength. For example, a monochromatic blue light
with a wavelength of 450 nm stimulates the red cones to a stimulus 0, green cones to
stimulus value 0 and blue cones to stimulus value 97. This set of ratios 0:0:97 will be
interpreted as blue by nervous system.

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Fig. 9. Absorption of spectra by cones

Source: ILLL in house

Value addition: Did you Know??

Heading text: Color Blindness

Body text: When a single group of color-receptive cones is missing from the eye,
the person is unable to distinguish some colors from others. This disorder is called
Color blindness. Depending on the type of cone missing, the color blindness can be
of following types:
1. Protanope: a person with loss of red cones.
2. Deuteranope: a person lacks green cones.
If either of these cones are missing, the person is unable to distinguish red, orange,
yellow and green color and is therefore called as red-green color blindness. This is a
sex-linked disorder that occurs exclusively in males because genes in the female X
chromosome code for respective cones. Red- green color blindness. red- green color
blindness is present predominantly in men, affecting 1 out of 12. Color blindness in
women is much more rare (1 out of 200) since males only have one X chromosome
(in females, a functional gene on only one of the two X chromosomes is sufficient to
yield the required photopigments).

At high light intensities, as in daylight vision, most people—92% of the male

population and over 99% of the female population – have normal color vision.
However, there are several types of defects in color vision that result from mutations
in the cone pigments. The most common form of colour blindness, red- green color
blindness. Colour blindness results through recessive mutation in one or more genes
encodes for the cone pigments. Genes encoding the red and green cone pigments
are positioned very close to each other on the X chromosome, whereas the gene
encoding the blue chromophore is located on chromosome 7.
Ishihara charts are used for determining color blindness.

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Fig. 10. Ishihara charts

Source: Textbook of Medical Physiology. Guyton and Hall

http://commons.wikimedia.org/wiki/File:Ishihara_compare_1.jpg
Image credit: This file is licensed under the Creative Commons Attribution-Share
Alike 3.0 Unported, 2.5 Generic, 2.0 Generic and 1.0 Generic license.

Processing and conduction of visual impulse

Role of various retinal cells in processing and conduction of visual impulse
1. Rods and cones transmit signals to the outer plexiform layer, where they synapse with
bipolar cells and horizontal cells.
2. Horizontal cells transmit signals horizontally in the outer plexiform layer from rods and
cones to bipolar cells.

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3. Bipolar cells transmits signals vertically from rods, cones and horizontal cells to inner
plexiform layer where they synapse with ganglion cells and amacrine cells.
4. Amacrine cells transmit signals in two directions, either directly from bipolar cells to
ganglion cells or horizontally to other amacrine cells or from axons of bipolar cells to
dendrites of ganglion cells in the inner plexiform layer..
5. Ganglion cells transmit output signals from retina through the optic nerve into the
brain.
6. Interplexiform cell transmits signals in the retrograde direction from the inner
plexiform layer to outer plexiform layer. This prevents lateral spread of visual signals by
horizontal cells in outer plexiform layer and also controls degree of contrast.

Visual Pathway for cones

Visual pathway for cones to ganglion cells is different from rods. The neurons and nerve
fibers that conduct visual signals for cones are larger than the ones for rods and hence the
transmission of the signal is two to five times faster than rods. The visual pathway for cones
is shown in the fig.

Neurotransmitters
Neurotransmitter glutamate is released by both rods and cones at their synapses with
bipolar cells.
Transmission of signals
The ganglion cells are the only retinal neuron that always transmits visual signals by action
potential rest of the retinal cells conduct by electrotonic conduction (direct flow of electric
current, not action potential from point of excitation to output synapses) to other cells of
the retina namely horizontal cells, amacrine cells, and ganglion cells. The ganglion cells pass
on the visual signals in the form of action potential to the neurons of lateral geniculate body
and soon after to the primary visual cortex. Visual image is decoded and interpreted in both
serial and parallel fashion.

Perception of vision
This is an intricate assimilation of senses of light, form, contrast and color of object. The
accessible visual field created in the retina and cortex is used to predetermine this visual
information of an object image.
1. Sense of light

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It is responsiveness to the light. The lowest brightness needed to induce a feeling of light is
termed as the light minimum. It can be determined when the eye is dark adapted for
minimum of 20-30 minutes. The visual adaptation is the adjustment that occurs within
the system of vision due to exposure to broad range of illumination for normal functioning of
eye. Two types of adaptation involved in this event:

 Dark adaptation (alteration in low light), and

 Light adaptation (alteration to high light).

Dark adaptation
It is the ability of the eye to adapt itself to decreasing illumination. If we go from sun-shine
into a dim light, we can only perceive the objects in dim light after many minutes have
passed. In course of this time interval, eye is adapting to low light. As described earlier,
vision can only be supplied by the rods in the darkened room. Therefore, rods are used
more in low level of light (i.e. in scotopic vision) and cones in high level of light (i.e. in
photopic vision).
If the person is fully dark adapted, his eye’s retina is many times (approx. one lakh
times) more responsive to brightness. If somebody has delayed dark adaptation, that
means he may has one of the diseases of rods, for example, retinitis pigmentosa and
vitamin A deficiency.

Light adaptation
When we go suddenly from a dark to a bright light or sun-shine, the light seems very
intense and still uncomfortably bright till the eyes are adapted to the higher level of light
and the visual threshold rises. This event in which retina adapts itself to bright illumination
is termed as light adaptation. Initially, the eye is extremely sensitive to light as rods are
overwhelmingly activated, and the visual image too bright and has poor contrast. However,
the rhodopsin, is soon inactivated (or bleached) through the high illumination- the rods
become unresponsive so that only the less- susceptible cones are operating and the
image becomes less and more clear.

Other mechanism of light and dark adaptation

In bright light, the circular muscle of the iris contract, causing constriction of the pupil
(decrease in the size). In the dim light, the radial muscles of the iris contract, causing an

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dilation of the pupil (increase in size) (Fig 11). Parasympathetic neurons stimulate the
circular muscle while sympathetic neurons stimulate the radial muscle of the iris.

Fig. 11: Autonomic Control of Pupillary Size

In bright, pupil constricts as circular muscles of iris contract and in dim light, pupil
dilates as radial muscles contract
Source: OpenStax College. "Autonomic Reflexes and Homeostasis." OpenStax- CNX. June 5,
2013. http://cnx.org/content/m46579/1.2/.
Image Credit: This file is licensed under the Creative Commons Attribution ShareAlike 3.0 License.

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.2. Sense of Form

It is the capability to distinguish between the shapes of the objects. Cones play a main role
in this process. So, the sense of object shape is most sharp at the fovea centralis, where
it has highest number of cones and reduces very quickly towards the periphery. In
medical observation, determination of the threshold of distinguishing two spatially apart
targets (a role of the fovea centralis) is called visual acuity.
Major differences exist between the peripheral retina and central retina. Near fovea, fewer
rods and cones converge on each on each optic fiber and the rods and cones also become
slender. These effects progressively increase the acuity of vision. At fovea centralis, there
are nearly 35000 slender cones and no rods are present and also nearly same number of
optic nerve leads from this part. Hence this region has high degree of visual acuity. The
peripheral retina on the other hand has much greater sensitivity to weak light.

Fig. 12. Difference in neural organization of retina at periphery and

at fovea
Source: ILLL in house

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 Physiology of Vision

3. Sense of Contrast
This is the capability of the eye to recognize minor changes in the luminance between areas
that are not separated by exact boundaries. Loss of contrast sensitivity causes mild
fogginess of the vision. Contrast sensitivity is affected by a variety of factors for example,
age, refractive errors, glaucoma, amblyopia, diabetes, optic nerve diseases and lenticular
changes. Additionally, contrast sensitivity may be weakening still in the presence of normal
visual acuity.

4. Sense of Colour
This is the ability of the eye to distinguish diverse colours transmitted through light of
different wavelengths. Colour vision results from different combinations of the three types
of cones (red, green and blue) and thus better experienced in photopic vision (Fig 13). In
low level of light (scotopic vision), you see all colours as grey and this phenomenon is
termed Purkinje shift.

Fig.13. Simulated appearance of a red geranium and foliage in

normal bright-light (photopic) vision, dusk (mesopic) vision,

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 Physiology of Vision

and night (scotopic) vision illustrate the Purkinje effect or

Purkinje shift
Source:http://en.wikipedia.org/wiki/Purkinje_effect#mediaviewer/File:Red_geranium_photo
ic_mesopic_scotopic.jpg

Value addition: Did you Know??

Heading text: How brain interprets a visual signal

―This figure shows how the brain uses mapping to make sense of visual information
from the eye. The green numbers in the figure correspond to the following steps:
Rays of light (blue) reflected off of an image are focused through the lens onto the
back of the eye, forming an upside-down image on the retina.
On the retina, those photocells that are hit by light from the image are activated.
These photocells are shown in white in this figure. Photocells that do not receive any
reflected light are not activated, and are shown in this figure. Thus, we can think of
the image as a pixellate map of activated and nonactivated photocells on the retina.
A nerve (gold) from each photocell connects to a particular location in the visual
cortex of the brain. The photocells that are activated (white) send a nerve impulse
to the brain, while the photocells that are not activated (black) do not send any
impulse to the brain. (Only a small sample of the nerves are shown in this figure.)
The brain, when it receives a collection of nerve signals from the eye, interprets
where each signal comes from, and reconstructs the pixellate map.

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 Physiology of Vision

The brain then interprets the pixellate map as an image.‖

Fig. 14. How brain interprets a visual signal

Source: http://www.chemistry.wustl.edu/~edudev/LabTutorials/Vision/Vision.html
Written for permission

Value addition: Video

Recap the concepts of vision physiology through this video:

<iframe width="420" height="315" src="//www.youtube.com/embed/_5dEO-LRV-g"

frameborder="0" allowfullscreen></iframe>

Source: You tube

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 Physiology of Vision

Summary

 Vision is the special sense of sight that is based on the transduction of light stimuli
received through the eyes.
 The wavelengths capable of stimulating the receptors of the eye-visible spectrum-
are between about 400 and 700nm. Different wavelengths of light within band are
perceived as different colors.
 Refraction, accommodation and changing of pupil size aids in achieving a clear image
of an object.
 Human eye is optically equivalent to a camera which has a lens system, a variable
aperture in form of pupil and a retina which corresponds to photographic film.
 About two third of the refractive power of the eye is provided by the cornea because
the refractive index of cornea is markedly different from the air whereas the
refractive indices of rest of lens system are not greatly different.
 The total refractive power of the internal lens of the eye is only 20 diopters, about
one third the total refractive power of eye
 The change in the shape of biconvex lens of human eye let the focusing power of
eyes adjust for far and close vision.
 When objects more than 6 metres away, the ciliary muscle of the ciliary body is
relaxed and lens is flatter because it is stretched in all directions. Stimulation of
parasympathetic nerves contracts both sets of ciliary muscles which relax the lens
ligament and hence lens becomes thicker and refractive power increased.
 With growing age, especially after 40, the lens grows larger and thicker and becomes
less elastic because of partial denaturation of lens proteins. Hence the ability to
change lens shape decreases and so is the power of accommodation.
 The major function of the iris is to increase or decrease the amount of light that
enters eye during darkness or brightness.
 With both the eyes open, the outer region of our total visual field is perceived by
only one eye known as zone of monocular vision. In the central portion, the fields
from the two eyes overlap known as the zone of binocular vision.
 Myopia, hyperopia and astigmatism are the refractive disorders of eye.
 An inverted image is formed on retina and the photoreceptor cells of retina finally
perceive the image to convey it to the CNS through following steps:
 Rhodopsin-retinal visual cycle and excitation of rods

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 Physiology of Vision

 Processing and conduction of visual sensation, performed through the

image processing cells of retina and visual pathway, and
 Perception of vision involves a role of visual cortex and associated regions of
cerebral cortex.
 The absorption characteristics of the pigments in the three types of cones help the
retina to differentiate the colors. The nervous system interprets the color on the
basis of ratio of stimulation by monochromatic light with a particular wavelength.
 Processing and conduction of visual impulse is carried out by various cells of retina:
rods, cones, bipolar cells, amacrine cells and ganglion cells which finally transfer the
signal to brain via optic nerve.
 Perception of vision is an intricate assimilation of senses of light, form, contrast and
color of object. The accessible visual field created in the retina and cortex is used to
predetermine this visual information of an object image.

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 Physiology of Vision

Exercises

A. Multiple Choice questions

1. The total refractive power of the internal lens of the eye is
A. 20 diopters
B. 6 diopters
C. 40 diopters
D. 60 diopters
2. The change in the shape of eye lens to let the focusing power of eyes adjust for far and
close vision is brought about by
A. Iris
B. Ciliary muscle
C. Extraocular muscle
D. Retina
3. The condition where each eye remains focused permanently at an almost constant
distance and can no longer accommodate for both near and distant vision is called
A. Emmetropia
B. Presbyopia
C. Hyperopia
D. Myopia
4. The condition where image is formed beyond retina for light rays coming from nearby
objects when all the ciliary muscles of eyes are relaxed is called
A. Emmetropia
B. Presbyopia
C. Hyperopia
D. Myopia
5. Photoactivation of electron in the retinal component leads to instantaneous conversion of
A. cis form of retinal to all-trans form.
B. cis form of retinol to all-trans form.
C. all-trans form of retinal to cis form.
D. all-trans form of retinol to cis form.
6. Excitation of rod causes increased negativity of intrarod membrane potential, which is a
state of
A. Hyperpolarization
B. Depolarization
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 Physiology of Vision

C. Hypopolarization
D. Neutralization
7. Three types of cones blue, green and red show peak absorbencies at light wavelengths of
A. 445, 535 and 570 nm
B. 535, 445 and 570 nm
C. 570, 535 and 445nm
D. 570, 445 and 535nm
8. Sense of object shape is most sharp at the fovea centralis, where it has highest number
of
A. Cones
B. Rods
C. Bipolar cells
D. Amarcrine cells
B. Short answer type questions
1. Define the following:
a. Rhodopsin
b. Visual acuity
c. Astigmatism
d. Accomodation
e. Presbyopia
2. Differentiate between:
a. Rods and cones
b. Monocular and Binocular vision
c. Myopia and hyperopia
d. Photopic and scotopic vision
C. Long answer type questions
3. Explain the neural pathways of vision.

4. Describe photochemical changes for initiation of vision.

5. What are differences between dark and light adaptation?

6. Explain the role of visible spectrum in colour vision.

7. Write short notes on the followings:

 Color vision

 Rhodopsin bleaching

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 Physiology of Vision

8. How and why the changes occur in size of pupil?

9. Explain accommodation for near vision and far vision.

10. Describe the refraction abnormalities and how they are corrected.

11. Explain the role of rod and cones and in vision.

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 Physiology of Vision

Glossary

Accommodation: It is the process to create a sharp image on the retina by changing the
thickness of lens as light rays from near objects must be bent more than those from distant
objects.
Amacrine cell: A type of cell in the retina that connects to the bipolar cells near the outer
synaptic layer and provides the basis for early image processing within the retina.
Binocular vision: The centre of the field of object view is binocular when it is seen by both
eyes that is termed as binocular vision.
Bipolar cell: Cell type in the retina that connects the photoreceptors to the retinal
ganglionic cells.
Colour vision: It is created by the three types of cones (red, green and blue) and so that
we can able to see colored object in bright light.
Cones: It is the photoreceptor which allow us see in brighter lights which stimulate cones to
produce colour vision.
Fovea centralis: It is the exact center of the retina at which visual stimuli are focused for
maximal acuity, where the retina is thinnest, at which there is nothing but photoreceptors.
Hyperopia: The condition where image is formed beyond retina for light rays coming from
nearby objects when all the ciliary muscles of eyes are relaxed.
Lens: The lens is a highly flexible, round and biconvex body, located just behind the pupil
and iris and helps focus images on the retina to facilitate clear vision.
Myopia: The condition where image is formed in front of retina for light rays coming from
distant objects when all the ciliary muscles of eyes are relaxed.
Photoreceptors: Photoreceptors are specialized cells in which light rays falls on objects are
transmitted to nerve impulses.
Phototransduction: It is a phenomenon of translation of light rays into nerve impulse is
known as phototransduction.
Presbyopia The condition where each eye remains focused permanently at an almost
constant distance and can no longer accommodate for both near and distant vision.
Refraction: Bending of light rays when it travelling through a medium in called refraction.
Rod: It is the photoreceptor which allow us see in dim light and we can see only black,
white, and all shades of gray in between.
Visual acuity: It is the property of vision related to the sharpness of focus, which varies in
relation to retinal position

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 Physiology of Vision

References

1. Tortora, G.J. & Grabowski, S. Principles of Anatomy & Physiology. 13th Edition,
p.642.
2. Moyes, C. D. and Schulte, P. M. (2006). Principles of Animal Physiology, p. 248.
3. Hill, R. W., Wyse, G. A. and Anderson, M. (2006). Animal Physiology. p.355.
4. Randall, D., Burggren W. and French, Kathleen (2001). Eckert Animal Physiology.
5. Widmaier, E.P., Raff, H. and Strang, K.T. (2008). Vander’s Human Physiology, XI
Edition, McGraw Hill.
6. Guyton, A.C. and Hall, J.E. (2011). Textbook of Medical Physiology, XII Edition,
Harcourt Asia Pvt. Ltd./W.B. Saunders Company.
7. Ganong, William F. Review of Medical Physiology. XXI Edition. Mc Graw Hill
8. Textbook of Physiology by Prof. A.K. Jain.
9. Anatomy And Physiology: In health and illness. Ross and Wilson (Tenth Edition)
10. Rushton, W. A. H. (1 June 1966). "Densitometry of pigments in rods and cones of
normal and color defective subjects" (PDF). Investigative Ophthalmology 5 (3): 233–
241. PMID 5296487.

Weblinks

 http://www.chm.bris.ac.uk/webprojects2003/rogers/998/Rhoeye.htm
 http://www.chemistry.wustl.edu/~edudev/LabTutorials/Vision/Vision.html
 http://webvision.med.utah.edu/book/part-ii-anatomy-and-physiology-of-the-
retina/photoreceptors/
 http://cnx.org/contents/b375ea7d-22d5-4f47-b10a-41dd93637896@4#fig-
ch14_01_14
 http://en.wikipedia.org/wiki/Photoreceptor_cell

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