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Olfactory Epithelium

located in the roof of the nasal cavity, contains olfactory receptor cells

2. to smell an odor it must be volatile and be dissolved in the fluid coating

the olfactory epithelium
3. the olfactory epithelium covers the superior nasal conchae on each side
of the nasal septum, and contains millions of bowling pin shaped
olfactory receptor cells
4. olfactory receptor cells are surrounded and cushioned by columnar
supporting cells
5. at the base of the olfactory epithelium lie the short basal cells.
6. olfactory receptor cells bipolar neurons, has thin apical dendrite with
olfactory cilia (increase receptive surface area) covered by mucus
(produced by supporting cells and olfactory glands)
7. mucus acts as solvent (dissolves airborne odorants)
8. unmyelinated axons of the olfactory receptor cells are gathered to the
filaments of the olfactory nerve (cranial nerve I)
9. olfactory nerve project through the openings in the cribriform plate of
the ethmoid bone, where they synapse in the overlying olfactory bulbs.
Specificity of the olfactory receptors

human can distinguish 10,000 or so odors, but research

suggests that our olfactory receptors are stimulated by different
combinations of olfactory qualities.

Activation of the olfactory receptors

odorant must be volatile

must dissolved in the fluid coating the olfactory epithelium

will stimulate the olfactory receptor by binding to protein

receptors in the olfactory cilium membranes and opening ion
sodium channels

this leads to a receptor action potential

Mechanism of smell transduction

odorant chemical binding to the G protein-associated receptor

sets the cAMP 2nd messenger system into motion,

which causes Na+ and Ca2+ channels to open.

Influx of Na+ causes depolarization and impulses transmission

The Olfactory Pathway

Action potentials of the olfactory nerve filaments are transmitted

to the olfactory bulb where the filaments synapse with mitral cells
(in glomeruli).

The mitral cells send impulses via the olfactory tract to the
olfactory cortex (smells will be interpret and identified)

Fibers carrying impulses from the olfactory receptors also project

to the limbic system (elicits emotional responses to odors).


Accessory structure of the eye

1. Eyebrows shade the eyes from sunlight, keep sweat out
2. Eyelids protect the eye from physical damage and from drying out
3. Conjunctiva a transparent mucous membrane that lines the eyelids
and anterior portion of sclera (the visible white part of the eye),
produces a lubricating mucous that prevent the eyes fro drying out
4. Lacrimal apparatus consists of lacrimal gland and ducts, the
secretion (tear) cleanses the eye, and drains into nasolacrimal duct
5. Six extrinsic eye muscles innervated by CN III (Oculomotor), IV
(Trochlear) & VI (Abducens), for moving the eyeball


Structure of the eyeball

1. Three tunics form the wall of the eyeball

a) Fibrous tunic the outmost layer, avascular, dense connective tissue,

includes 2 regions: cornea (anterior portion) sclera (posterior
sclera protects the eyes and gives it shape
cornea allows light to enter the eye
b) Vascular tunic (uvea) middle layer, vascular, includes 3 regions:

choroid (posterior portion) - contains pigment which helps

absorb light and prevent the light from scattering and
reflecting within the eye; provides nutrients to the eye.


ciliary body (anterior portion) ciliary muscles controls lens



iris (the most anterior portion) - the visible colored part of

the eye, controls pupil size

c) Sensory tunic (retina) innermost layer, vascular, include 2 layers

1) outer - pigmented layer absorbs light and prevent lights from
scattering, act as phagocytes and store vit.A.
2) inner neural layer. From posterior to anterior, the neural
layer is composed of three types of neurons :
photoreceptors (rods and cones), bipolar cells and
ganglion cells. The other 2 types of neuron; horizontal
cells and amacrine cells plays a role in visual
processing. Ganglion cell axons form the optic nerve,
which exits via the optic disc (blind spot)

2. Lens : avascular, biconcave, transparent, flexible, its shape can be
changed to allow focusing of light on retina. It is enclosed in a thin, elastic
capsule and held in place just posterior to the iris by suspensory ligament.
3. Internal chambers and fluids
a) Posterior cavity filled with vitreous humor. Vitreous humor:
(1) transmit light,
(2) supports the posterior surface of the lens and holds the neural retina
firmly against the pigmented layer,
(3) contributes to intraocular pressure, helping to counteract the pulling
force of the extrinsic eye muscles.
b) Anterior cavity filled with aqueous humor, for transporting nutrients & O 2
to lens and cornea and it carries away their metabolic wastes

Is a thin protective covering of epithelial cells. It protects the

Conjunctiva cornea against damage by friction (tears from the tear glands help
this process by lubricating the surface of the conjunctiva)

Is the transparent, curved front of the eye which helps to converge

the light rays which enter the eye
Is an opaque, fibrous, protective outer structure. It is soft


connective tissue, and the spherical shape of the eye is maintained

by the pressure of the liquid inside. It provides attachment
surfaces for eye muscles
Has a network of blood vessels to supply nutrients to the cells and


remove waste products. It is pigmented that makes the retina

appear black, thus preventing reflection of light within the eyeball.
Has suspensory ligaments that hold the lens in place. It secretes

Ciliary body

the aqueous humour, and contains ciliary muscles that enable the
lens to change shape, during accommodation (focusing on near
and distant objects)
Is a pigmented muscular structure consisting of an inner ring of
circular muscle and an outer layer of radial muscle. Its function is


to help control the amount of light entering the eye so that:

- too much light does not enter the eye which would damage
the retina
- enough light enters to allow a person to see
Is a hole in the middle of the iris where light is allowed to continue


its passage. In bright light it is constricted and in dim light it is



Is a transparent, flexible, curved structure. Its function is to focus

incoming light rays onto the retina using its refractive properties
Is a layer of sensory neurones, the key structures being
photoreceptors (rod and cone cells) which respond to light.
Contains relay neurones and sensory neurones that pass impulses

along the optic nerve to the part of the brain that controls vision

A part of the retina that is directly opposite the pupil and contains


only cone cells. It is responsible for good visual acuity (good



Blind spot

Is where the bundle of sensory fibres form the optic nerve; it

contains no light-sensitive receptors
Is a transparent, jelly-like mass located behind the lens. It acts as


a suspension for the lens so that the delicate lens is not damaged.


It helps to maintain the shape of the posterior chamber of the



Helps to maintain the shape of the anterior chamber of the eyeball


The focusing of the light rays onto the retina from objects at different
distances is known as accommodation.

Focusing involves the bending or refraction of light rays

Viewing a distant object

light rays entering the eye from distant objects are virtually parallel and
require little refraction (bending) to bring them to a focus.

The circular ciliary muscles relax, forming a wider ring and this causes
them to pull on the suspensory ligaments, making them tight.

The ligaments in turn pull on the lens, stretching it and making it flatter.
This less rounded (less convex) lens causes less refraction.

Accommodation to long distance is aided by the internal pressure of the

eye caused by the vitreous humour. This helps to widen the ciliary
muscles and tighten the ligaments.

Viewing a near object

If we now change to looking at a near object, the light rays entering the
eye from the object are spreading out (diverging) and require more
refraction to bring them to a focus

The circular ciliary muscles contract, forming a narrower ring.

The suspensory ligaments become loose as they are not being pulled by
the muscles.

The lens is not stretched and because it is elastic it reverts to its natural
rounded shape.

This fatter (more convex) lens causes more refraction, allowing a sharp
focus to be achieved.


Myopia is commonly known as short-sightedness.

It means that the eye is unable to focus on distant objects, making them
seem blurred.

Vision can usually be corrected with concave glasses or contact lenses,

or, in some cases, with laser eye surgery.

How the eye focuses

Light coming into the eye is focused onto the retina - an area on the rear
wall of the eyeball.

Seventy per cent of light entering the eye is focused by the cornea (a clear
dome which forms the outer covering of the pupil). The rest is focused by
the lens, which sits behind the cornea.

If the light is focused properly on the retina, we see a clear image.

If the light focuses in front of the retina, the image is fuzzy. See the
illustration above.

With myopia, the cornea is too curved or the eyeball too long. This means
that images are focused in front of the retina rather than directly on it.


Hyperopia is a defect of vision caused by an imperfection in the eye

(often when the eyeball is too short or when the lens cannot become
round enough), causing inability to focus on near objects, and in extreme
cases causing a sufferer to be unable to focus on objects at any distance.

As an object moves towards the eye, the eye must increase its power to
keep the image on the retina.

If the power of the cornea and lens is insufficient, as in hyperopia, the

image will appear blurred.

Minor amounts of hyperopia are sometimes left uncorrected, however,

larger amounts may be corrected with convex lenses in eyeglasses or
contact lenses. Hyperopia is sometimes correctable with various refractive
surgery procedures.

Photoreception is the process by which the eye detects light energy.
To approach this subject several areas need to be discussed

The functional anatomy of photoreceptor cells.

The location and packaging of the visual pigments that absorb the light

The chemistry of the visual pigments and their response to light.


Activation of the photoreceptors and their response to varying levels of

light intensity


The photoreceptor cells are found in the retina at the back of the eye.

There are 2 types of photoreceptor cells

1) rods
2) cones


There are present in large very large numbers, with approximately 120
million rods and 6 millions cones per eye.

Rods and cones have three main regions. The outer segment is the light
sensitive region containing photosensitive pigments. Here light energy is
converted into a generator potential.

In rods this region contains up to 1000 membrane-lined vesicles. The

photosensitive pigment rhodopsin is embedded in the membranes of the

Vesicles are also present in cones but they consist of infoldings of the
outer membrane. The cones have a different photosensitive pigment,
iodopsin, embedded in the membranes.

The inner segment of both rods and cones contains many mitochondria to
provide the energy to resynthesise the visual pigments after they have
been broken down by light.

It also contains the nucleus and many ribosomes to synthesis proteins

such as those needed to make the vesicles.

The synaptic zone is the equivalent of the synaptic bulb in neurons and
allows the information of synapses with bipolar cells which in turn link to
the optic nerve and then to the brain.


Visual Transduction

Visual transduction is the process by which light initiates a nerve

impulse. The structure of a rod cell is:

The detection of light is carried out on the membrane disks in the outer
segment. These disks contain thousands of molecules of rhodopsin, the
photoreceptor molecule.

Rhodopsin consists of a membrane-bound protein called opsin and a

covalently-bound prosthetic group called retinal.

Retinal is made from vitamin A, and a dietary deficiency in this vitamin

causes night-blindness (poor vision in dim light).

Retinal is the light-sensitive part, and it can exists in 2 forms: a cis form
and a trans form:

In the dark retinal is in the cis form, but when it absorbs a photon of light it
quickly switches to the trans form.

This changes its shape and therefore the shape of the opsin protein as
well. This process is called bleaching.

The reverse reaction (trans to cis retinal) requires an enzyme reaction and
is very slow, taking a few minutes.


This explains why you are initially blind when you walk from sunlight to a
dark room: in the light almost all your retinal was in the trans form, and it
takes some time to form enough cis retinal to respond to the light indoors.

The final result of the bleaching of the rhodopsin in a rod cell is a nerve
impulse through a sensory neurone in the optic nerve to the brain.

However the details of the process are complicated and unexpected. Rod
cell membranes contain a special sodium channel that is controlled by

Rhodopsin with cis retinal opens it (sodium channel) and rhodopsin with
trans retinal closes it.

This means in the dark the channel is open, allowing sodium ions to flow
in and causing the rod cell to be depolarised.

This in turn means that rod cells release neurotransmitter in the dark.

However the synapse with the bipolar cell is an inhibitory synapse, so

the neurotransmitter stops the bipolar cell making a nerve impulse.

In the light everything is reversed, and the bipolar cell is depolarised and
forms a nerve impulse, which is passed to the ganglion cell and to the


Rods and Cones

Why are there two types of photoreceptor cell? The rods and cones serve two
different functions as shown in this table:
Outer segment is rod shaped

Outer segment is cone shaped

106 cells per eye, found mainly in the

109 cells per eye, distributed throughout
fovea, so can only detect images in
the retina, so used for peripheral vision.
centre of retina.
Good sensitivity can detect a single
photon of light, so are used for night

Poor sensitivity need bright light, so

only work in the day.

Only 1 type, so only monochromatic


3 types (red green and blue), so are

responsible for colour vision.

Many rods usually connected to one

bipolar cell, so poor acuity (i.e. rods
are not good at resolving fine detail).

Each cone usually connected to one

bipolar cell, so good acuity (i.e. cones
are used for resolving fine detail such
as reading).


Visual Acuity
Visual acuity the amount of detail that can be seen. The cones are responsible
for high visual acuity (high resolution). Although there are far more rods than
cones, we use cones most of the time because they have fine discrimination and
can resolve colours. To do this we constantly move our eyes so that images are
focused on the small area of the retina called the fovea. You can only read one
word of a book at a time, but your eyes move so quickly that it appears that you
can see much more. Spatially, much more clarity is perceived in cones than for
the rods. This is because one cone cell synapses to one bipolar cell which in
turn synapses onto one ganglion cell as the information is relayed to the visual
cortex. The more densely-packed the cone cells, the better the visual acuity. In
the fovea of human eyes there are 160 000 cones per mm 2, while hawks have 1
million cones per mm2, so they really do have far better acuity.


A. Outer (external) ear
1. auricle (pinna) - composed of elastic cartilage covered with thin skin and
an occasional hair
helix (the rim) - thicker
lobule (earlobe) fleshy, lacks supporting cartilage
f(x): auricle to direct sound waves into external auditory canal.
2. external auditory canal (meatus)

short curved tube that extends from the auricle to the eardrum

the entire canal lined with hairy skin, sebaceous gland and
ceruminous glands.

Ceruminous glands secrete cerumen (earwax) to provides sticky

trap for foreign particles

Tympanic membrane (eardrum) thin, translucent, connective

tissue membrane, covered by skin (externally) and by mucosa
(internally). It is a boundary between outer and middle ear.

Eardrum vibrates when sound waves reach it.


A. Middle ear (also called tympanic cavity)

1. filled with air
1. separated from outer ear by tympanic membrane (eardrum), transfers
vibration (generated by sound wave) to ossicles

(1) malleus or hammer - a tiny bone that passes vibrations

from the eardrum to the anvil.
(2) incus or anvil - a tiny bone that passes vibrations from
the hammer to the stirrup.
(3) stapes or stirrup - a tiny, U-shaped bone that passes
vibrations from the stirrup to the cochlea. This is the
smallest bone in the human body (it is 0.25 to 0.33 cm

2. three ossicles transfer vibration from eardrum to oval window, which sets
the fluids of inner ear into motion, and stimulates hearing receptors
oval window - An oval opening located on the medial wall of the tympanic
cavity, leading into the vestibule, to which the base of the stapes is
connected and through which the ossicles of the ear transmit the sound
vibrations to the cochlea.
3. communicates with nasopharynx through pharyngotympanic auditory
tube (Eustachian tube) for balancing air pressure
Eustachian tube - a tube that connects the middle ear to the back of the
nose; it equalizes the pressure between the middle ear and the air outside.
When you "pop" your ears as you change altitude (going up a mountain or
in an airplane), you are equalizing the air pressure in your middle ear.
4. round window - An opening on the medial wall of the middle ear that
leads into the cochlea and is covered by the secondary tympanic


B. Inner (internal) ear

1. Bony labyrinth (osseous) is filled with perilymph
a) Vestibule is the central egg shaped cavity of the bony labyrinth
b) cochlea - a spiral-shaped, fluid-filled inner ear structure; it is
lined with cilia (tiny hairs) that move when vibrated and cause a
nerve impulse to form by the Organ of Corti (the receptor organ
for hearing).
c) semicircular canals - three loops of fluid-filled tubes that are
attached to the cochlea in the inner ear. They help us maintain
our sense of balance.
2. Membranous labyrinth is surrounded by the perilymph, its interior
contains endolymph

perilymph and endolymph these two fluids conduct the sound

vibrations involved in hearing and respond to the mechanical forces
occurring during changes in body position and acceleration.

cochlear nerves - these carry electro-chemical signals from the inner ear (the
cochlea) to the brain.



Sound is the compression and rarefaction of air, or, in other words, alternating air
pressure. The distance between the pressure peaks is called the wavelength.
The frequency of the sound is {wavelength -1 x speed of sound} . The frequency
determines the pitch of the sound. Humans can detect sound in the frequency
range 20 to 20,000Hz (Hz = cycles, or waves, per second).
Sound - Vibrations transmitted through an elastic material or a solid, liquid, or
gas, with frequencies in the range of 20 to 20,000 hertz, capable of being
detected by human organs of hearing.
The numbers in the diagram below indicate the sequence of events in the
detection and transduction of sound waves.

1. Sound waves enter the external ear and are directed towards the
tympanic membrane.


2. Air molecules under pressure cause the tympanic to vibrate. Low

frequency sound waves produce slow vibrations and high frequency
sounds produce rapid vibrations. These move the malleus on the other
side of the membrane.
3. The handle of the malleus strikes the incus causing it to vibrate.
4. The vibrating incus moves the stapes in and out and vibrates the oval
window. The total force of the sound wave is transferred to the oval
window, but, because the oval window is much smaller the force per unit
area is increased 15-20 times. Additional mechanical advantage is gained
from the leverage in the middle ear bones. This is necessary because the
fluid in the inner ear is more difficult to move than air and thus sound must
be amplified.
5. The sound waves that reach the inner ear through the oval window set up
pressure changes that vibrate the perilymph in the scala vestibuli.

6. Vibrations in the perilymph are transmitted across the vestibular

membrane to the endolymph of the cochlear duct, and also up the scala
vestibuli and down the scala tympani. The vibrations are transmitted to the
basilar membrane which in turn vibrates at a particular frequency,
depending upon the position along its length {high frequencies vibrate the


window end where the basilar membrane is narrow and thick, and low
frequencies vibrate the apical end where the membrane is wide and thin}.

7. The cilia of the hair cells, which contact the overlying tectorial membrane,
bend as the basilar membrane vibrates, this opens ion channels and
causes the entry of ions into the hair cell and a generator potential
develops. If large enough, the generator potential causes transmitter
release from the hair cells which excites the afferent nerve. Displacement
of the stereocilia in the direction of the tallest stereocilia (called the
kinocilium in hair cells of the vestibular system and immature auditory
system) is excitatory and in the opposite direction is inhibitory. One theory
suggests a mechanical link to ion channels which opens a "trap door" as it
is pulled taut. This is a little fanciful and it is thought that the ion channels
are located at the base of the stereocilia but are indeed
mechanoreceptors, in that they respond to mechanical pressure. The
endolymph surrounding a hair cell is K+ rich and so K+ (and calcium)
enter the hair cell, causing a depolarisation.
8. The action potentials are transmitted along the cochlear branch of the
vestibulocochlear nerve, activating auditory pathways in the central
nervous system, eventually terminating in the auditory area of the
temporal lobe of the cerebral cortex.
9. Finally, the vibrations in the scala tympani are dissipated out of the round
window, into the middle ear.



Afferent nerves from the cochlear (spiral) ganglion terminate in the cochlear
nucleus in the brainstem.
Axons from neurons in the cochlear nucleus project to the superior olive, inferior
colliculus, medial geniculate nucleus (of the thalamus) and the auditory cortex
(Brodman areas 41 and 42).
Note that auditory input projects to both sides of the cortex. The superior olive
and the inferior colliculus send efferent fibres back to the stapedius and tensor
tympani muscles respectively. These muscles are concerned with protecting the
middle ear bones from overload.