Dr Aziz Zarina, Assistant Professor

Special senses

The special senses are the senses that have specialized organs devoted to them:

 vision (the eye)

 hearing and balance (the ear, which includes the auditory system and vestibular system)
 smell (the nose)
 taste (the tongue)

Visual system

Visual perception is the ability to interpret the surrounding environment using light in the
visible spectrum reflected by the objects in the environment. The resulting perception is also
known as visual perception, eyesight, sight, or vision. The various physiological components
involved in vision are referred to collectively as the visual system. The visual system in animals
allows individuals to assimilate information from their surroundings.
The eyes are the sensory organs that are extension of the brain. The eyes receive visual
information from the environment and transmit them to the visual sensory area of the brain for
interpretation.

The pineal and parapineal glands are photoreceptive in non-mammalian vertebrates, but
not in mammals. Birds have photoactive cerebrospinal fluid (CSF)-contacting neurons within the
paraventricular organ that respond to light in the absence of input from the eyes or
neurotransmitters.

Accessory Structure of the Eye

Eyelids - (Palpebrae)
The free margins of the eyelids form the palpebral fissure. The edges where the upper and
lower eyelids meet are called the lateral and medial canthus. The palpebral margins are
lined by eyelashes. The eyelids are supported internally by sheets of connective tissue
that form the tarsal plate.
Glands associated with the eyelids include:
- Glands of Zeis - These are sebaceous glands associated with the eyelashes.
- Tarsal glands - (Meibomian glands) - These glands line the inner margin of the
lid and produce a lipid-rich product that prevents the lids from sticking together.
- Lacrimal caruncle - This is a mound of tissue in the medial canthus that produces
thick secretions.
The conjunctiva
It is the epithelium that lines the inner surface of the eyelids and continues onto the outer
surface of the eye. The palpebral conjunctiva lines the inner surface of the eyelid and the
bulbar conjunctiva lines the anterior surface of the eye. The conjunctiva consists of a
specialized stratified squamous epithelium that is thick until it lines the transparent
cornea where it is relatively thin.

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Lacrimal Apparatus
The lacrimal apparatus produces, distributes and removes tears. Tears are continually
produced and reduce friction, remove debris, prevent bacterial infection and provide
nutrients and oxygen for the conjunctiva. Lacrimal glands are located near the lateral
canthus of the eye, it produces tear in response to parasympathetic nerve stimulation.
Tears flow over the cornea and are drained into the nose by the nasolacrimal duct.
Nictitating membrane or the third eyelid is found in the medial canthus and it aids in
protection of the eye and its glands also produce tears. Tear is also produced by the
harderian glands, which is one of the lacrimal glands found in many bird and mammals
except in carnivores.

Structure of eye

The eye is a ball-shaped organ which is placed in the orbital fossa. The eyeball is hollow and is
divided into two cavities:
a. Posterior cavity
This cavity takes up most of the internal space and extends anteriorly to the lens. It
contains a gelatinous vitreous body and is also called the vitreous chamber.
b. Anterior cavity
This cavity is in the front of the lens and is filled with a clear liquid called aqueous
humor. It is further subdivided into anterior and posterior chambers by the iris.

The wall of the eye has three concentric layers called tunics

1. Outer layer (fibrous tunic): The eyeball is enveloped by a tough outer protective fibrous
sheet of connective tissue called sclera. Anteriorly the sclera is modified into a stratified
squamous epithelial layer called the cornea, which is transparent. It provides mechanical
support and protection of the eye. In birds it plays a role in accommodation.

2. Middle vascular layer (uvea): It has three regions namely

a. Choroid: Sclera is posteriorly lined by highly vascular and deeply pigmented layer
(melanocytes) called choroid. It serves a nutritive function for ocular tissue. Melanocytes
prevent light that escapes past the retina from being reflected back into the retina where it
would blur the image. In nocturnal animals this layer contains a reflecting pigment called
tapetum lucidum which causes “eye shine” in the night when light enters the eye. The
tapetum lucidum allows the retina to make optimal use of available light but visual acuity is
reduced.

b. Ciliary body: It is a thickened extension of choroid. It forms a muscular ring around the lens
and supports iris and lens. It secretes fluid called aqueous humor

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c. Iris: It is a diaphragm of varying size and is a pigmented structure containing dilator and
constrictor smooth muscle which controls diameter of its central opening called pupil. It has
two pigmented layers:
- Posterior pigment epithelium: Block stray light from reaching retina
- Anterior border layer: contain pigmented cells called chromatophore. Higher
concentration of melanin in chromatophore gives iris black, brown or hazel colour. If
scanty melanin is present, light will reflect from posterior pigment layer giving a
blue, green or grey colour to the iris.

Lens (aquula):
The lens is a transparent, biconvex structure in the eye that, along with the cornea,
helps to refract light to be focused on the retina. The lens is more flat on its anterior side
than on its posterior side. The lens, by changing shape, functions to change the focal
distance of the eye so that it can focus on objects at various distances, thus allowing a
sharp real image of the object of interest to be formed on the retina. This adjustment of
the lens is known as accommodation. It is the primary structure responsible for
accommodation. It is made up of an elastic lens capsule containing a jelly-like substance.
It is suspended behind pupil by a ring of fibers called suspensory ligaments which are
attached to ciliary body. The convexity of the lens can be altered by the ciliary muscles of
the ciliary body. This helps in focusing images of objects at varying distance from the eye
onto the retina. Ciliary
muscle contraction
increases the convexity
of the lens and focuses
near objects on the
retina. When lens
becomes opaque it is
called cataract. In
humans, the refractive
power of the lens in its
natural environment is
approximately
18 diopters, roughly one-
third of the eye's total
power. The lens itself
lacks nerves, blood vessels, or connective tissue.

The space between the cornea and the lens is called the anterior chamber and the
posterior chamber lies between the iris and the suspensory ligament. Both these chambers
are filled with a clear water-like fluid called aqueous humor. The ciliary process of the

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 ciliary body in the posterior chamber produce aqueous humor and it flows into the
anterior chamber through the pupil. It is absorbed into the venous system at right angle
between the cornea and the iris. If the absorption is obstructed, intraocular pressure
increases which is called glaucoma.

Behind the lens is a chamber filled with gelatinous fluid called the vitreous
humor. The vitreous humor is a hydrogel that contains hyaluronic acid and collagen
fibers. It supplies nutrition to the retina. Behind the vitreous humor is the neural retina.
The retina is interrupted at a point called optic disk, where the axons of the retina’s
ganglion cell layer leave the retina to the brain. The optic disk gives rise to the optic
nerve (2nd cranial nerve). Arteries and veins enter the eye at the optic disk and provide
nutrition to the retina. The choroid blood vessels provide the remaining nutrition to the
retina.
3. Inner Layer (neural tunic) / Retina

It is the innermost layer of the eye which is light sensitive. Its main functions are
photoreception and impulse transmission. The retina is considered part of the central nervous
system (CNS) and is actually brain tissue. It is the only part of the CNS that can be
visualized non-invasively. It is pressed smoothly against rear of eye ball by pressure of vitreous
humor and is attached to eye at two parts; optic disk and anterior margin. Retina can be detached
by blow to the head or insufficient pressure from vitreous humor. A detached retina causes blurry
area in the field of vision and may lead to blindness if it remains separated for too long from
choroid.

The retina consists of two layers:

a. Outer Pigmented layer / Retinal Pigmented Epithelium (RPE):

It is the posterior most layer of retina adjacent to the choroid. It is a single cell layer of
darkly pigmented cuboidal cells whose basal process interdigitate with receptor cells of
the retina. It protects and nourishes the retina, stores Vitamin A, removes waste products,
prevents new blood vessel growth into the retinal layer and absorbs light not absorbed by
the photoreceptor cells; these actions prevent the scattering of the light and enhance
clarity of vision.

b. Inner Neural layer

The neural layer consists of three layers of cells that include the photoreceptors, cells that
initiate the processing of visual information and blood vessels that supply the neural
layer. The neural layer extends anteriorly to the boundary called the ora serrata.

The layers of the neural layer from the outermost layer to the innermost are:
i. Photoreceptors
There are two types of photoreceptors:
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 - Rods are more sensitive to light but there is only one type of rod and color
discrimination is not possible with rods.
- Cones are less sensitive to light but there are three types of cones with
sensitivities in different regions of the light spectrum. Cones provide color
discrimination and greater detail.

ii. Bipolar cells

The photoreceptors synapse with bipolar cells. They exist
between photoreceptors and ganglion cells. they are the first order neurons of the
visual pathway. They act, directly or indirectly, to transmit signals from the
photoreceptors to the ganglion cells. They can synapse with photoreceptors and
they also accept synapses from horizontal cells. The transmission of visual
information through the bipolar cells is modulated by horizontal cells found in
this layer The bipolar cells then transmit the signals from the photoreceptors or
the horizontal cells, and pass it on to the ganglion cells directly or indirectly
(via amacrine cells).
Horizontal cells: They are the laterally interconnecting
neurons that connect different photoreceptors. They may also synapse with
bipolar cells. They help integrate and regulate the input from
multiple photoreceptor cells.

iii. Ganglion cells

They are the largest neurons of the retina, arranged in a single layer close
to the vitreous body. They are second –order neurons of visual pathway. Most of
the ganglion cells receive input from multiple bipolar cells. The ganglion cell

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 axons form the optic nerve. The transmission at this layer is modulated by
amacrine cells.
Amacrine cells: They are inhibitory neurons in the retina and
project their dendritic arbors interacting with retinal ganglion
cells and/or bipolar cells. amacrine cells affect the output from bipolar
cells.

There are no photoreceptors present in the retina where the optic nerve leaves the eye, and this
region is called blind spot

Photoreceptors

They are the only neurons that are directly sensitive to light. They are cells that absorb
light and generate a chemical or electrical signal. There are two kinds of photoreceptors; rods
and cones which produce visual image. They are named for the general shape of cells. Rod cells
are responsible for night (scotopic) vision ie they cannot distinguish colours from each other.
Cones function in bright light, they are responsible for day (photopic) vision as well as colour
vision. Rods and cones are derived from the stem cells that produce ependymal cells of the brain.

Structure of rods and cones

Each rod or cone has an outer segment that points towards the wall of the eye and an
inner segment facing the interior. These two
segments are separated by a narrow constriction
containing nine pairs of microtubules

Outer segment of rods and cones

It is actually highly modified cilium to absorb light.

In rods, the outer segment is cylindrical and
resembles a stack of coins in a paper roll; there is
plasma membrane around the outside and a neatly
arranged stack of about 1000 membranous discs
inside. Each disc is densely studded with globular
proteins called visual pigment rhodopsin. The
membranes hold these pigment molecules in a
position that results in most efficient light
absorption.

A cone has outer segment similar to rods except

that the outer segment tapers to point and the discs
are not detached from the plasma membrane but

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are parallel infoldings of it. 40% of the entire mass of outer segment is contributed by visual
pigments

Inner segment:

The inner segment contains mitochondria and other organelles. At its base it gives rise to cell
body. Cell body contains the nucleus and process that synapse with retinal neurons in the next
layer.

Visual pigment

In rods, the visual pigment is called rhodopsin or visual purple. Each molecule consists
of a protein called opsin and a vitamin A derivative called retinal (retinene). Opsin is embedded
in the disc membrane of rods outer segment. All rods contain a single kind of rhodopsin with an
absorption peak at a wavelength of 500 nm. The rods are less sensitive to light of other
wavelengths. Rhodopsin has a low threshold of excitability and is easily stimulated by low-
intensity light. Rod cells are about 300 times more sensitive to light than are cones.

In cones, the pigment is called photopsin (iodopsin). Retinal moiety is same as that of
rhodopsin but opsin moieties have different amino acid sequences that determine which
wavelengths of light the pigment absorbs. Based on this, there are three kinds of cones, which
are identical in appearance but optimally absorb different wavelength of light. They are
- Blue cone (430 nm)
- Green cone (535 nm)
- Red cone (575 nm)

Photochemical iodopsin requires relatively high-intensity light to be stimulated, thus cones

have higher threshold of excitability than rods.

Young – Helmholtz trichrome theory

It states that colour is a function of three different visual pigments, each being associated
with different cones. The visual pigments have a particular absorption spectrum with a
absorption maximum wavelength in red, green and blue. Therefore, each cone responds
maximally to wavelengths corresponding to absorption maximum of its pigment. There is an
overlap of spectra to allow the perception of other colours. Colour vision requires the presence of
atleast two visual pigments of overlapping sensitivity to luminance and with overlapping light
absorption spectra. This was first elucidated as the Young-Helmholtz trichrome theory.

When the light intensity decreases to a point where it is too weak to stimulate cone cells,
colour vision disappears. This is why night and twilight scenes appear gray.

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 Perception of white light: About equal stimulation of all the red, green and blue cones
gives one the sensation of seeing white. There is no wavelength of light corresponding to white,
instead white is a combination of all the wavelengths of the spectrum.

Colour blindness : when a single group of colour receptive cones is missing from the eye, the
person is unable to distinguish some colours from others. If either of blue or green cone is
missing, the person is unable to distinguish red from green and therefore is said to have red-
green colour blindness. The person with loss of red cones is called protanope. A colour blind
person who lacks green cones is called deuteranope. Red-green colour blindness is a genetic
disease that occurs almost exclusively in males but is transmitted through female.

Different animals have different kinds of color vision. Some have very poor color vision
and others have very good color vision. In fact some birds and bees have super color vision and
see colors that humans don't see. Dogs, cats, swine, mice, rats and rabbits have very poor color
vision. In fact, they see mostly greys and some blues and yellows. Monkeys, ground squirrels,
birds, insects, and many fish can see a fairly good range of color. Cattle, sheep and goat are red-
green colour blind. Domestic animals have dichromatic color vision.

Distribution of photoreceptors in the retina

The human retina contains about 120 million rod cells, and 6 million cone cells. The
number and ratio of rods to cones varies among species, dependent on whether an animal is
primarily diurnal or nocturnal. Light rays from the cornea travel through the unmyelinated,
transparent ganglion cells and bipolar cells before reaching the rods and cones. These cause
some distortion to the light rays. The fovea is an area located at the back of the retina, dorso-
lateral to the optic disc, where the light rays are not distorted because ganglion cells and bipolar
cells are pushed apart so that light falls directly on the photoreceptor. In man and primates cones
are concentrated at fovea which provides sharp visual image. In domestic animals fovea is
absent, but a homologous area is present referred to as area centralis or macula or visual streak.
Retina of domestic animals contains mostly rods while retina of birds contains mostly cones. At
the origin of optic disc no photoreceptors are present and this area is called blind spot.

The photochemical reaction / bleaching reaction / rhodopsin-retinal visual cycle

The receptor potential of rods and cones is generated by reactions that are initiated by
light being absorbed by chemicals called visual pigments. The reactions are called bleaching and
are cyclical. A metabolic pathway restores the original visual pigment. While bleached, the
visual pigment cannot respond to additional light stimuli. Each visual pigment absorbs only some
of the wavelength of light.

A good example of bleaching reaction is the bleaching of rhodopsin. Rhodopsin contains

a protein opsin and the carotenoid pigment retinal. The retinal present is 11-cis retinal.

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 When light energy is absorbed by rhodopsin the rhodopsin begins within trillionth of a
second to decompose. The cause of this is photoactivation of electrons in the retinal portion of
the rhodopsin, which leads to an instantaneous change of the cis-form of retinal to all-trans form,
which still has the same chemical structure as the cis-form but has a different physical structure
(a straight molecule rather than an angulated molecule). Because the three-dimensional
orientation of the reactive sites of all-trans retinal no longer fits with the orientation of the rective
sites on the protein opsin, it begins to pull away from opsin.

The intermediate reactions for conversion from 11-cis to all-trans is

- Rhodopsin is first converted to bathorhodopsin, which is a partially split combination of

the all-trans retinal and opsin
- Bathorhodopsin itself is an extremely unstable compound and decays in nanosecond to
lumirhodopsin
- Lumirhodopsin then decays in microseconds to metarhodopsin I and then in about a
millisecond to metarhodopsin II which is called activated rhodopsin that produce
electrical changes in the rods that then transmit the visual image to CNS
- Metarhodopsin II much more slowly (in seconds) separates completely into all-trans
retinal and opsin

Reformation of rhodopsin

When trans- retinal dissociates from opsin, it is transported to the pigment epithelium,
converted to cis-retinal, returned to the rod outer segment and reunited with opsin. This process
requires metabolic energy and is catalyzed by enzyme retinal isomerase. Once 11-cis retinal is
formed, it automatically recombines with opsin to form rhodopsin, which then remains stable
until its decomposition is again triggered by absorption of light energy.

There is a second chemical route by which all-trans retinal can be converted to 11-cis
retinal. This by conversion of all-trans retinal first into all-trans retinol, which is one form of
vitamin A. Then, all-trans retinol is converted into 11-cis retinol under the influence of enzyme
isomerase. And finally, the 11-cis retinol is converted to 11-cis retinal that combines with opsin
to form rhodopsin.

Vitamin A is present both in the cytoplasm of the rods and in the pigment layer of the
retina. Therefore vitamin A is normally always available to form new retinal when needed. On
the other hand, when there is excess retinal in the retina, the excess is converted back to vitamin
A, thus reducing the amount of light sensitive pigment in the retina. Night blindness
(nictalopia) occurs in any person with severe vitamin A deficiency. The reason is that not
enough vitamin A is available to form adequate quantities of retinal. Therefore the amount of
rhodopsin that can be formed becomes severely depressed. This condition is called night
blindness because the amount of light available at night is too little to permit adequate vision,

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although in daylight the cones can still be excited despite reduction of their colour pigments as
well.

It takes about 5 minutes to regenerate 50% of bleached rhodopsin. Cone cells are less
dependent on the pigment epithelium and regenerate half their pigment in about 90 seconds.

Dark and light adaptation

Light adaptation
This occurs when we move from the dark into bright light. The bright light momentarily
dazzles us and all we see is white light because the sensitivity of the receptors is set to dim
light. Rods and cones are both stimulated and large amounts of the photopigment are broken
down instantaneously, producing a flood of signals resulting in the glare.
Adaption occurs in two ways:
- The sensitivity of the retina decreases dramatically.
- Retinal neurons undergo rapid adaptation inhibiting rod function and favouring the cone
system.

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 Within about one minute the cones are sufficiently excited by the bright light to take
over. Visual accuracy and colour vision continue to improve over the next ten minutes. During
light adaptation retinal sensitivity is lost.

Dark adaptation

Dark adaptation is essentially the reverse of light adaptation. It occurs when going from
a well light area to a dark area. Initially blackness is seen because our cones cease functioning in
low intensity light. Also, all the rod pigments have been bleached out due to the bright light and
the rods are initially nonfunctional. Once in the dark, rhodopsin regenerates and the sensitivity
of the retina increases over time (this can take approximately one hour). During this adaptation
process reflexive changes occur in the pupil size.

Phototransduction of visual signal / Mechanism of vision / Generating optic nerve signal

The photoreceptors have cGMP-gated sodium channels in the

outer segment, whereas non gated potassium channels and Na+-
K+ ATPase pumps are present in the inner segment membrane.
Potassium leaks out through nongated potassium channels. Na-
K pump maintain intracellular concentrations of Na & K.
Voltage gated calcium channels are present at the synaptic
terminal portion of the photoreceptors.

In darkness, the photoreceptors have high level of cGMP.

cGMP binds to the sodium channels and opens the channels,
triggering sodium influx. This generates inward current called
dark current. The photoreceptors are depolarized during
darkness (-40mV). This causes opening of voltage gated
calcium channels at the synaptic terminal and inflow of calcium.
This helps in release of neurotransmitter glutamate at synaptic
sites on bipolar and horizontal cells of retina. Glutamate, though
usually excitatory, functions here as an inhibitory
neurotransmitter and will cause IPSP (inhibitory post synaptic
potential) at the bipolar cells.

On exposure to light, the concentration of cGMP is decreased through a series of biochemical

steps triggered by photopigment activation.

- A light photon interacts with the retinal in a photoreceptor cell. The retinal
undergoes isomerisation, changing from the 11-cis to all-trans configuration
- Retinal no longer fits into the opsin binding site.

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- Opsin therefore undergoes a conformational change to metarhodopsin II.
- Metarhodopsin II is unstable and splits, yielding opsin and all-trans retinal.
- The opsin activates the regulatory protein transducin.
- Transducin activates phosphodiesterase or PDE.
- PDE breaks down cGMP to 5'-GMP. This lowers the concentration of cGMP and
therefore the sodium channels close.
- Closure of the sodium channels causes hyperpolarization (-80mV) of the cell due to the
ongoing efflux of potassium ions.
- Hyperpolarization of the cell causes voltage-gated calcium channels to close.
- As the calcium level in the photoreceptor cell drops, the amount of the neurotransmitter
glutamate that is released by the cell also drops.
- A decrease in the amount of glutamate released by the photoreceptors causes excitation
of bipolar cells.
- Bipolar cells show graded potential change. If it is of sufficient magnitude cause
excitation of ganglion cells
- Ganglion cells develop action potential which is carried to the cerebral cortex for further
processing in the form of optic signals

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Visual pathway

The hyperpolarization of rods and cones alters the activity of bipolar cells, which in turn
influences the impulse frequency in the ganglion cells. The horizontal cells influence the bipolar
cells and the amacrine cells influence the ganglion cells to detect contrast and contour.

The ganglion cell axons from the temporal retina travel along the optic nerve to the optic
chiasma (where the right and left optic nerves cross) and then pass ipsilaterally to the lateral
geniculate nucleus of the thalamus on the same side of brain

The ganglion cell axons from the nasal retina come to the optic chiasma and cross the
contralateral geniculate nucleus of the thalamus.

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 The cells of the lateral geniculate nucleus project to the visual cortex of the occipital lobe
of cerebral cortex.

Image arising from the left field of vision is perceived in

the right visual cortex and from right lateral field of vision in the
left visual cortex.

A part of the optic nerve will terminate in the superior

colliculi of midbrain to initiate visual reflex and pretectal nucleus
of brain stem for pupillary constriction and a part of optic nerve
will terminate in the suprachiasmatic nucleus of hypothalamus to
set circadian rhythms.

Visual acuity (VA)

It commonly refers to the clarity and sharpness of vision. The

highest acuity will be achieved if each optic fiber carries signal
from a single photoreceptor cell. Cone cells provide better acuity
than rod cells. This is because hundreds of rod cells feed signals
via bipolar cells to a single ganglion cell, whereas only a few cone cells feed signals to a single
ganglion cell. The highest visual acuity is provided by area centralis where the cone cells
concentration is more.

Types of vision

Monocular vision/Periscopic vision

Animals with laterally placed eyes, view the objects with one eye at a time,
independently of the other eye. This is due to wider visual angle between optic axis and median
line of the eye. Example; Amphibians, reptiles, most birds (Owls are exceptions)

Binocular vision

Primates, carnivores and some birds have the power of converging of the eyes, thus view
the same object simultaneously with both eyes. This is due to parallel optic axis and median line,
which provide overlap of field of vision.

Stereo vision

Primates, cats and other felines have three dimensional view because of small angle
between optic axis and median line of the eye. The left and the right eye show dissimilarity in
viewing an object and these two retinal images are fused in the visual center of the brain giving
information about height, width as well as depths of the object; the three dimensional picture.

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 Accommodation of eye

It is the ability of the eye lens to adjust its focal length by changing its curvature for
focusing the image to the retina for both near and distant vision.

For near vision the anterior curvature of the eye lens is increased by the action of the
ciliary muscles. The ciliary body reduces the tension in the suspensory ligament, causes bulging
of the anterior curvature of eye lens towards the less resistant anterior chamber of eye. On the
other hand, relaxation of ciliary muscles creates tension in the suspensory ligament, cause
flattening of the eye lens and this helps in distant vision.

In horse, the retina is placed at different distances from cornea because of the ramp
shaped retina. The ciliary muscles are weak and are unable to alter the curvature of the lens to
change the focal length. The animal simply raises or lowers its head or eye to allow the image to
fall on the retina.

In domestic animals – the predatory animals as dogs and cats can increase the power of their
lenses more than horses and ruminants ( ie, accommodative power is more in dogs and cats). Birds are
good accommodators and each eye in birds can accommodate independently of the other. In birds ability
to accommodate reduces with age (as human).

Presbyopia

It is the condition of the eye characterized by gradual loss of power of accommodation,

which is associated with loss of emmetropia (ability to focus parallel rays of light on retina).
This is due to decreased elasticity of the eye lens with increasing age.

Refraction errors of eye

If the eye lens is unable to adjust its focusing capacity to focus image on the retina, it is
called ametropia. It is of three types

Myopia / short or near vision

Abnormal increase in the anterior posterior diameter of the eyeball or the excessive
refraction of the eye lens causes focusing of the image just before the retina. It can be corrected
by concave lens which reduce the refractive power of the eye lens thus focus image exactly on
the retina.

Hypermetropia (long or far sight)

It is a common defect in wild animals, shortening of the anterior posterior diameter of the
eyeball or less refraction of the eye lens, thus image fall behind the retina. It can be corrected by
convex lens which increase the refractive power of the eye lens thus focus image exactly on the
retina.

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Astigmatism

The difference in the radius of curvature of the cornea causes the light to focus not to a
common point on the retina, which results in both myotropic and hypermetropic visions. This
defect can be corrected by cylindrical lens.

Visual peculiarities in birds

The ostrich has the largest eye of any land vertebrate, with an axial length of 50 mm, twice that
of the human eye. Kingfishers and Hummingbirds have two foveae. Convergence (many photoreceptors
synapse with one bipolar cell) is very low in birds and hence have high resolution for their vision around
eight times greater than that of humans especially in hawks and eagles.

In diurnal birds, 80% of the receptors may be cones (90% in some swifts) whereas nocturnal owls
have almost all rods and in the rods itself 2 types of visual pigments. Most birds are tetrachromatic,
possessing four types of cone cells each with a distinctive maximal absorption peak and pigeons are
pentachromatic.

The pecten occuli is a structure seen only in birds, having folded tissue which projects from the
retina. It is well supplied with blood vessels and appears to keep the retina supplied with nutrients, and
may also shade the retina from dazzling light or aid in detecting moving objects. Pecten oculi is
abundantly filled with melanin granules which have been proposed to absorb stray light entering the bird
eye to reduce background glare.

Electroretinogram (ERG)

The electroretinogram (ERG) is recording of the electrical activity generated by neural

and non-neuronal cells in the retina in response to a light stimulus. It is composed principally of
A, B and C waves, although A and B waves are normally used clinically.

A wave : It corresponds primarily to activation of visual

receptor cells and bleaching of visual pigment

B wave: It corresponds primarily to the activation of the

bipolar cell layer of the retina. The wave is from ganglion
cells, amacrine cells and bipolar cells.

C wave: It is wave from pigment epithelium

Scotopic ERG is recorded after the eye is dark

adapted (kept in darkness for 20-30 minutes). It represent the activation of both rods and cones
and therefore the wave will be higher in amplitude when compared to the photopic ERG that is
recorded in light adapted eye where the recording will be only from cones (because all rhodopsin
will be in bleached position). Recording of both types of ERG allows detection of night
blindness and abnormalities of the light adapted eye.
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Electroocculogram: It is a record of potential difference between the front and back of the eye
ball. The recording electrode is placed subcutaneously at the lateral canthus of the eye, and the
reference electrode is placed at the forehead. The potential difference varies with eye movement
and therefore eye movement can be recorded with this.

Tonometry is the procedure to determine the intraocular pressure (IOP), the fluid pressure inside
the eye. The normal range is 10-21 mmHg. It is an important test in the evaluation of patients at
risk from glaucoma. Most tonometers are calibrated to measure pressure in millimeters of
mercury.

Ophthalmoscopy, also called funduscopy, is a test that allows to see inside the fundus of the
eye and other structures using an ophthalmoscope (or funduscope). It is done as part of an eye
examination and may be done as part of a routine physical examination. It is crucial in
determining the health of the retina, optic disc and vitreous humor.

The Auditory System

The auditory system is designed to detect and analyse sound in the environment. Much of
animal communication relies on this system. Sounds are travelling vibrations through air or
liquid characterized by alternating regions of increased pressure caused by compression of
molecules and region of decreased pressure caused by rarefaction of molecules. The auditory
system perceives the frequency of sounds as pitch (unit is hertz) and their amplitude as loudness
(unit is decibel). The neural perception of sound is called hearing. Hearing requires at least
one ear but localization of sound requires two ears as the auditory system must detect the
difference in time of arrival or intensity of sound impinging on the two ears. Animal’s sense of
hearing is enhanced by their ability to move their ears to scan of different sounds and locate
where the sound is coming from.

Structure of ear

The ear consists of three parts; external ear (outer ear), middle ear and inner ear.

External ear: The external ear is made up of pinna (skin covered auricular cartilage) and
external auditory meatus (ear canal).

Pinna: It guides sound waves down the ear canal and acts as a resonating structure. Orientation
of pinna toward a sound source increases sound detection. In birds, external ear lacks a flap or
pinna and is hidden by specialized feathers known as auricular feathers. These feathers protect
the opening, reflect sound and enhance sound wave collection.

17
The auditory canal conducts the sound waves into the middle ear. It is lined by waxy secretion
called as cerumen. This filters the air entering the ear to avoid infections. There is a separation
between the outer ear and the middle ear called as the tympanic membrane or commonly called
as the ear drum. The ear drum transfers the sound waves to the vibrations which are conducted
towards the inner ear by the middle ear. In dogs, the ear canal is long and bent and often traps
wax or provides an ideal habitat for mites, yeast and bacteria.

Middle ear: It is air filled cavity, separated from the external ear by the tympanic membrane and
from the inner ear by vestibular window (oval window) and cochlear window (round window).
containing ossicles and Eustachian tube.

The ear ossicles are bony structures namely malleus (hammer), incus (anvil), and stapes
(stirrup). They conduct the vibrations toward the inner ear. The malleus is attached to the
tympanic membrane while the foot plate of the stapes fit into the vestibular window in the bony
wall between middle and inner ear. The ossicles provide a mechanical linkage between the
tympanic membrane and the oval window. Thus vibration of tympanic membrane reaches the
inner ear. In birds only one bone called columella is present so transmission of sound is less
efficient.

Eustachian tube (auditory tube): It is a collapsible tube that connects middle ear with
oropharynx. It opens periodically to equalize middle ear pressure with external atmospheric
pressure and to clear fluid from middle ear. In horse, it has a ventral diverticulum called guttural
pouch.

Two skeletal muscles (tensor tympani and stapedius muscle) are located in the middle ear which
alters the transmission of vibration between the ear drum and oval window.

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Inner Ear (Labyrinth)

It is made up of bony labyrinth, within it is located the membranous labyrinth. The

membranous labyrinth is surrounded by fluid called perilymph. Membranous labyrinth consists
of cochlea (acoustic part) and vestibular organ (non-acoustic part).

Cochlea :

Cochlea looks like a coiled up snail shell. It consists of three parallel, fluid filled canals
that are coiled around a central bony axis called modiolus. The number of cochlear turns varies
between 2 and 4 in different mammalian species. There is no clear relationship between the
number of turns and hearing ability. Modiolus contain ganglion cells of auditory portion of
cranial nerve VIII called spiral ganglion.

The fluid filled tubes are

Scala vestibuli (vestibular duct): It is the uppermost canal that starts at the oval window and ends
at the tip (apex) of the cochlea. It is separated from scala media by Reissner’s membrane
(vestibular membrane).

Scala tympani
(tympanic duct): It is
the lowermost canal
that starts at the round
window and
terminates at the apex
of cochlea and is
separated from the
scala media by the
basilar membrane.

Scala media (cochlear

duct): It is the fluid
filled canal present
between the scala
vestibuli and scala tympani.

Scala vestibuli and scala tympani are connected with each other at the apex of cochlea at
a region called helicotrema. Both the scala vestibuli and scala tympani are filled with a fluid
called perilymph, while scala media is filled with endolymph. The perilymph has a composition
similar to common interstial fluid. The endolymph has a much higher potassium concentration
and a positive electric potential (+80mV) relative to interstitial fluid.
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Organ of corti (spiral organ):

The sensory organ of the auditory system is the organ of corti. It is the transducer that
converts vibrations into nerve impulses. The organ of corti sits atop the basilar membrane in
scala media which is narrow and stiff at the base near the oval and round windows and broad and
elastic at the apex. The organ of corti consists of sensory cells, supporting cells and tectorial
membrane.

The sensory cells are called hair cells, because they have thin, stiff sensory hairs
(microvilli or stereocilia) protruding into the endolymph. The stereocilia are arranged in
increasing order of height
from one end of hair cell to
the other end. Stereocilia are
connected by tip links at their
tips. Tip link is composed of
filamentous material, which
connect stretch sensitive ion
+
channel (K channel) at the tip of one stereocilium with the side of adjacent longer stereocilium.
The cells are stimulated by shear movements of the sensory hairs. The hair cells are located in 4-
6 rows from the oval window to the tip of cochlea. The hair cells in the row nearest the axis of
the cochlea are called inner hair cells, while the hair cells in the remaining 3-5 rows are called
outer hair cells. The inner hair cells detect sound while the outer hair cells amplify sound waves
(cochlear tuning). The inner hair cells do not have their own axons but form synapses with
sensory nerve fibers. The cells of the sensory nerve fibers are located in a ganglion in the bony
axis of the cochlea. The nerve fibers extend from the spiral ganglion to the brain in cranial nerve
VIII (vestibulocochlear nerve). 90-95% are axons originating from synapse with inner hair cells.

Surrounding the hair cells is a highly organized system of supporting cells and directly
above is a gelatinous membrane called the tectorial membrane which is secreted by the

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supporting cells. It projects as a horizontal shelf from the canal wall closest to the axis of the
cochlea. The stereocilia of the inner hair cells just touch the tectorial membrane while the
stereocilia of the outer hair cells are embedded in the tectorial membrane.

Mechanism of hearing

Sound waves enter the ear, vibrate the tympanic membrane causing the middle ear ossicles to
vibrate. Since the surface area of the tympanic membrane is large compared to the attachment of the
middle bones, it concentrates the sound energy allowing it to vibrate the bones of the middle ear. Without
this focusing, vibration of the structures would be greatly reduced. This function of causing air vibration
to cause vibration of a solid and eventually a liquid is referred to as impedance matching. The stapes is in
contact with the oval window causing the vibration
to be transmitted to the fluid of the the scala
vestibuli. The moving fluid can either transfer the
movement laterally to the other 2 tubes, the scala
media and scala tympani or travel to the end of the
scala vestibuli and, through an opening called the
helicotrema into the scala tympani. These
travelling waves in turn cause vibrations in the
basilar membrane, which causes tectorial
membrane to slide back and forth over inner and
outer hair cells of the organ of Corti. The up and
down movement of the basilar membrane causes
bending of stereocilia. Bending of stereocilia in the
direction of tallest stereocilia cause pulling of tip
link. Pulling of tip link opens the ion channel
present at the tip of stereocilia. This triggers K+ ion
influx into the hair cells which will depolarize the
hair cells. Depolarization causes opening of
voltage-gated calcium channels at the base of the
cell and subsequent influx of calcium. This will
cause release of neurotransmitter into the synaptic
cleft between the hair cells and terminal ends of
cochlear nerve (spiral ganglion). An action
potential develops in the cochlear nerve.

The bending of stereocilia away from the tallest stereocilium causes hyperpolarization of hair
cells.

Hair cells at different locations along the basilar membrane respond to different frequencies of
sound waves. High frequency (short wavelength) detected at the base and low frequency (high
wavelength) detected at apex.

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Auditory pathway

The nerve impulse from the cochlear nerve to

carried to the vestibulocochlear nerve. The
vestibulococlear nerve terminates in cochlear nucleus
of medulla. The axons from the cochlear nucleus
terminates into inferior colliculus of mid brain. From
here the axons will project into the median geniculate
body of thalamus and from there to the primary and
secondary auditory cortex of temporal lobe of cerebral
cortex. Primary auditory cortex functions to study
features of sound (frequency, loudness and pattern).
The secondary auditory cortex process the information
further for recognition of sound.

The range of frequencies that can be detected varies

among the species. Humans can detect sounds in the
range from 20 to 20,000 Hz, although sensitivity is
poor at both extremes. Some animals can perceive
frequencies much higher than 20,000 Hz. The upper
pitch unit of dogs is generally thought to be about
twice that of humans. The silent dog whistle emits
some frequencies too high for humans to hear but below the upper pitch limit for dogs.
Elephants can perceive sound below 14-24 Hz through vibrations of feet.

The cochlea of the bird differs considerably from that of the mammal. It is short and
nearly straight, rather than coiled. Birds generally have a narrower range of hearing than
mammals, probably because of mechanical constraints on the transmission of sound to the
cochlea. Some birds can hear ultrasonic sounds.

Auditory agnosia: Inability to hear

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 Vestibular system

Sensory information about motion,

equilibrium, and spatial orientation is provided
by the vestibular apparatus, which in each ear
includes the anterior utricle, posterior saccule,
and three semicircular canals. The otolith
organs (utricle and saccule) detect gravity
(information in a vertical orientation) and
linear movement. The semicircular canals
detect rotational movement.

Otolith organs (gravireceptor)

Each of these chambers has a 2 by 3 mm patch called macula composed of hair cells and
supporting cells. Each hair cell of a macula has 40-70 stereocilia and one true cilium called
a kinocilium. The tips of these cilia are embedded in an otolithic membrane (gelatinous
membrane). This membrane is
weighted down with protein-
calcium carbonate granules
called otoliths. These otoliths add
to the weight and inertia of the
membrane and enhance the sense
of gravity and motion. With the
head erect, the otolithic membrane
bears directly down on the hair
cells and stimulation is minimal.
When the head is tilted, however,
the otolithic membrane sags and
bends the stereocilia, stimulating the hair cells. Any orientation of the head causes a combination
of stimulation to the utricles and saccules of the two ears. Movement of head causes bending of
stereocilia which will cause opening of K+ channel leading to depolarization of the hair cells.
This will cause release of neurotransmitter into the synaptic cleft between the hair cells and
terminal ends of vestibular nerve. An action potential develops in the vestibular nerve.

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Semicircular ducts

Rotational acceleration is detected by the three semicircular ducts, each housed in an osseous
semicircular canal of temporal bone. The anterior and posterior semicircular ducts are positioned
vertically, at right angles to each other. The lateral semicircular duct is about 30oC from the
horizontal plane. The orientation of the duct causes a different duct to be stimulated by rotation
of the head in different planes. The semicircular ducts are filled with endolymph. Each duct
opens into the utricle and has a dilated sac at one end called an ampulla. Within the ampulla is a
mound of hair cells and supporting cells called crista ampullaris. The hair cells have stereocilia
and a kinocilium embedded in the capula (a gelatinous membrane that extends from the crista to

the roof of ampulla. When the head turns the duct rotates, but the endolymph lags behind. It
pushes capula, bends the stereocilia and stimulates hair cells. After 25 to 30 seconds of continual
rotation, the endolymph catches up with the movement of duct and stimulation of the hair cells
ceases.

Vestibular pathway

The information from the vestibular hair cell is conveyed through the vestibular nerve to the
vestibulocochlear nerve. The nerve fibers from vestibulocochlear nerve projects ipsilaterally
vestibular nuclei. The vestibular nuclei on either sides of the brain stem exchange signals
regarding movement and body position. These signals are sent down the following projection
pathways.

- To the cerebellum. Signals sent to the cerebellum are relayed back as muscle movements
of the head, eyes, and posture.
- To nuclei of cranial nerves III, IV, and VI. Signals sent to these nerves cause the
vestibulo-ocular reflex. They allow for the eyes to fix on a moving object while staying in
focus.

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 - To the reticular formation. Signals sent to the reticular formation signal the new posture
the body has taken on, and how to adjust circulation and breathing due to body position.
- To the spinal cord. Signals sent to the spinal cord allow quick reflex reactions to both the
limbs and trunk to regain balance.
- To the thalamus. Signals sent to the thalamus allow for head and body motor control as
well as being conscious of body position.

Vestibulo-ocular reflex (VOR) is a reflex eye movement that stabilizes images on

the retina during head movement by producing an eye movement in the direction opposite to
head movement, thus preserving the image on the center of the visual field. It is by means of
vestibular projections from the semicircular ducts.

Olfactory system

The olfactory system, or sense of smell, is the part of the sensory system used for
smelling (olfaction). The senses of smell and taste (gustatory system) are often referred to
together as the chemosensory system, because they both give the brain information about the
chemical composition of objects through a process called transduction. Most mammals and
reptiles have a main olfactory system (in the dorso-caudal part of nasal cavity) and an accessory
olfactory system (Vomeronasal or Jacobson’s organ near the external nares). The main olfactory
system detects airborne substances (volatile and water soluble chemicals), while the accessory
system senses fluid-phase stimuli.
Olfaction is essential for the localization of food, reflex stimulated secretion of digestive
enzymes, and detection of danger. The olfactory cells are part of the specialized olfactory
epithelium found on the ethmo-turbinate bones of the nasal cavity. The olfactory mucosa
occupies a relatively large area in dogs (100 cm2) compared with that in humans (about 5 cm2).
The olfactory cells give rise to the olfactory nerve fibers that terminate in the olfactory bulb.

Mechanism of olfaction
- Olfaction starts when the odorant enters the nasal cavity and comes into contact with the
mucus layer that lines the olfactory epithelium of the nose.
- This mucus contains odorant binding
proteins which allow lipophilic odorants to
dissolve in the aqueous mucus layer.
- The olfactory receptor cells are
bipolar neurons where one end of the
neuron ends in the olfactory epithelium and
the other ends as a synaptic terminal with
the neurons of the olfactory bulb of the
brain.

25
 - The olfactory receptor cells are covered by cilia which project into the nasal
passage. These are non-motile, and contain odorant receptor proteins.
- The odorant receptor proteins are G-proteins. These proteins are of multigene family
and contain many numbers of genes coding for odorant receptors.
- Each receptor expresses single receptor protein. Thereby the ability to distinguish even
the slightest change in the odorants.
- As the odorant molecules bind with the
receptor, the receptor undergoes conformational
change that sends signals to the G-protein
associated with it. The activated protein act
through the adenylyl cyclase pathway and cause
generator potential.
- If the depolarization is large enough, then
the action potential is triggered which travel to
the bipolar neuron.
- The action potential is transmitted from
the bipolar neuron to the synaptic terminal with
the neurons of the olfactory bulb and they carry
the information to the olfactory bulb.
- The olfactory bulb is a complex neural structure. It has a lining of ball-like neural
junction known as glomeruli. Each glomerulus receives signals only from a single type
of receptor cell. Thereby, these glomeruli act as smell files to sort out the type of smell
of the odorant.
- The fibers leaving the olfactory bulb pass through two routes.

 Subcortical route: It goes to the regions of the limbic system mainly to the
medial sides of the temporal lobe which is concerned with olfaction and known as
primary olfactory cortex. It includes hypothalamic involvement and coordinates
smell with behavioral reactions.
 Thalamic-cortical route: This route is important for conscious perception and
fine discrimination of smell.

- The odorants are cleared from the nasal passage very quickly and the olfactory system
adapts quickly, so the sensitivity to the smell of a particular odorant diminishes after a
short exposure to the smell. The odorants are cleared by enzymes named as odorant-
clearing enzymes. They resemble the detoxification enzymes in liver.

Accessory olfactory system (Vomeronasal or Jacobson’s organ)

- Absent in humans and apes
- Located along anterior part of nasal septum

26
 - Open at one end and forms blind sac at other end
- Location of opening is variable (In mouse opens into nasal cavity while in cow opens into
oral cavity)
- Receptor similar to the main olfactory system
- Sensitive to large, non-volatile molecules that cannot reach the main olfactory system
- Has got a pumping mechanism that serves to suck molecules (pheromones) into the organ

Variation in olfaction
Animals differ in their ability to detect odours. Animals like dogs, are very sensitive in
detection of odour and are called as macrosmatic; those that can detect odour but with less
sensitivity like birds are termed microsmatic; those that lack olfactory apparatus like dolphins
and whales are termed anosmic.

Ungulates and rhesus monkeys detect the female in heat by the odour of the vaginal
secretions. The olfactory system involved in maternal recognition, regulation of reproduction,
mother-young interaction, establishment of dominance.

Gustatory system (sense of taste)

All mammals and birds have taste organs called taste buds that are found on the tongue, roof of
mouth, soft palate and pharynx. There are almost 15,000
taste buds in the porcine oral cavity and throat, while 1
550 in the lizards, but only 24 in chickens. They are
onion shaped structure that contains numerous taste
receptor cells. It has a pore that opens onto the surface
where the tastants (chemicals from the food) enter and
contact the receptors. Each bud has about 40-50
elongated receptor cell and sustentacular or supporting
cells. The receptor cells are arranged around taste pore.
Each receptor cell has several microvilli or taste hairs on its apical region that extend into the
taste pore. The basal surface of the receptor cells are innervated by terminal gustatory nerves.
Each taste receptor cell expresses more than one taste receptor protein. The taste receptors are
not neurons instead they are epithelial cells which secrete neurotransmitter onto a primary
afferent neuron. Single taste neurons synapse with more than one taste receptor cells. Olfaction
and gustation work together closely and that is the reason why the taste perception is dependent
on smell perception of an item.

There are primarily five tastes like salty, sour, sweet, bitter, and umami.

27
Salty: The salt taste is mainly due to the Na+ ions and metal ions. So the salty food causes
opening of the Na+ channels. The substance namely Amiloride inhibits the receptor channels and
diminishes the salt sensation of the receptors.

Sour: The sour taste is due to the H+ ions in the food. The sour perception is done via the
Na+ channels, which are permeable to H+ ions, but compete with them and hence it is done only
in species which have less Na+ ions in their saliva. In other animals, it is by acid-sensing ion
channels. It causes opening of the Na+ channels in response to the change in pH

Sweet: The sweet perception is by the binding of the sweet substance to the G-protein which
activates G-protein gustducin, which signals the adenylate cyclase pathway. This taste is
triggered by organic compounds like sugar, alcohol, aldehyde, ketones

Umami:- Taste is triggered by amino acid such as aspartic acid and glutamate in meat. This taste
is also evoked by inosinic acid and qunilic acid. The glutamate bind to a G protein- coupled
receptor and activate a second messenger system.

Bitter taste: - These elicited mainly alkaloids, long chain organic compounds with nitrogen and
many toxic substances. In bitter signaling pathway, the G protein Gustidin activate the second
messenger pathway.

Gustatory pathway

Taste impulses from the anterior two third of the tongue pass through chorda tympani branch of
the facial nerve‚ then to nuclei tractus
solitarius of the brain stem, while the
posterior one-third sends the taste
sensations through the glossopharyngeal
nerve to nuclei tractus solitarius. The
tractus solitaries will send signal to
 Superior and inferior salivary
nuclei of brain stem to regulate secretion
of salivary glands
 Ventrobasal region of thalamus

Vagus receives the sensory inputs from

the base of the tongue and other parts of
the pharyngeal region and send signal to
the ventobasal region of thalamus.

The signal from the ventobasal region of thalamus is carried to the parietal lobe of cerebral
cortex for processing.

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 Sense

Any of the faculties by which man and animals become aware of stimuli originating inside or
outside the body. Eg. smell, sight, hearing. Modality is the distinct property of each sensation.

Classification of senses

1. Special senses: eg. Vision, hearing, smell, etc.

2. General senses

- Cutaneous senses: Are those with receptors on the skin, eg. Touch, pressure, warmth,
cold etc.
- Visceral senses: are those concerned with perception of internal environment

Receptors
Receptors are specialized neuronal structures that detect diverse stimuli from the internal
and external environment

Classification

1. Depending on the position and function,

a. Teloceptors / Teloreceptors : (Distant receivers) which are concerned with events at a
distance. Eg. rods and cones, hair cells.
b. Exteroceptors: concerned with external environment at hand. Eg. Receptors for touch,
cold, warmth, etc.
c. Interoceptors: Concerned with internal environment. Eg. Baroceptors in carotid and
aortic bodies, osmo receptors in blood vessels.
d. Proprioceptors: which provide information about the position of the body in space at
any given instant, eg. Muscle spindle, Golgi tendon organ.

II. Depending on their terminations

a. Encapsulated nerve endings
(i) End bulb/Kraisen bulb - (for cold)
They are oval or spherical in shape. They are seen in lips, mucous membrane of
tongue, cheek, soft palate, epiglottis, nasal cavity etc. They are receptors for cold.
(ii) Meissners Corpuscle – (for touch)
They are oval or elongated structures. They are seen especially in hairless portions of
skin such as palm, fingertip, lips etc. They are receptors for touch
(iii) Pacinian corpuscle - (For deep pressure and vibration)
It is elliptical in shape. They are present in deeper subcutaneous connective tissue of
hand, feet, peritonium etc. They are receptors for deep pressure.

29
 b. Free nerve endings

They are seen throughout the body. They detect pain (nociceptors), cold or touch and
subconscious stimuli for reflex action.

III. According to energy sensitivity

a. Mechanoceptors – that respond to mechanical energy. Eg. pacinian corpuscles,

merkel’s discs in skin ( provide information on pressure, position, and deep static touch
features such as shapes and edges), ruffni endings (respond to sustained pressure).
b. Chemoceptors - that respond to chemical energy. Eg. chemoceptors in aortic and
carotid body.
c. Thermoceptors – that respond to temperature. eg. kraisen bulb.
d. Nociceptors - pain; physical or chemical damage
e. Photoreceptors - That respond to electromagnetic radiation like light; rods & cones of
the eye

Proprioceptors.
The two important proprioceptors are Muscle spindle and Golgi tendon organ

Muscle spindle: Distributed throughout the belly of the muscle and send information about the
length of muscle or rate of change of its length and they transmit information to spinal cord,
cerebellum and cerebral cortex and help in controlling muscle contraction.

The muscle spindle receptors can be excited in two ways.

- Lengthening the whole muscle will obviously stretch the mid portion of the spindle and
therefore excite the receptor.
- Contraction of the end portion of the muscle also stretches the mid portion and excites the
receptor.
Golgi tendon organ: Is an encapsulated sensory receptor through which a small bundle of
tendon fibers pass; immediately before their point of fusion with the bone. The golgi tendon
organ is stimulated by the tension produced by the small bundle of muscle fibers.

The major functional difference between the muscle spindle and golgi tendon organ is
that the spindle detects the muscle length while the tendon organ detects the muscle tension.

Receptor function

Whatever the type of stimulus that excites the receptor, its immediate effect is to change
its membrane potential. This change in potential is called receptor potential. The receptor
potential is produced by change in the receptor membrane permeability to produce a potential
difference. When this receptor potential rises above the threshold for it produces action potential

30
in a nerve fiber attached to the receptor. The more the receptor potential rises above the threshold
value the greater will be the frequency of action potential.

Maximum amplitude of receptor potential is 100 mV ie when the membrane becomes maximally
permeable to Na+.

Properties of Receptors

1. Specificity: Receptors are specific that they will respond to only one kind of stimuli.

Eg: Photoreceptors respond to light stimuli alone.

2. Relation with strength of stimuli: Receptor will respond only if the stimuli are of threshold
strength. Increase in strength of stimuli beyond the threshold strength will not produce
additional response.

3. Projection: When any part of the sensory pathway is stimulated conscious sensation is
referred to the location of the receptor and this is called law of projection.

4. Adaptation: The perception of a sensation may disappear if the stimulus is applied

continuously. The time of adaptation vary with the receptors. Mechanoceptors adapt
more rapidly when compared to other receptors. Some of the chemoceptors and pain
receptors never adapt completely. But pacinian corpuscle adapt with in fraction of a sec.

5. After images: Sensation persists even though the stimulus has been removed.
Eg: Looking at bright light, then closing the eye the sensation of seeing the bright light is
left.

Expanded tip Free nerve endings Krause corpsucle Meissners corpuscle

receptor

31
Pacinian corpuscle Ruffinis end organ Tactile hair Markel's disc

32