The Special Senses

Objectives

 At the end of this chapter the students will;

1. Explain physiology of olfaction
2. Discuss physiology of gustation
3. Enumerate physiology of vision
4. List physiology of hearing
5. Discuss physiology of equilibrium

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 Outlines

 Introduction
 Olfaction

 Gustation

 Vision

 Hearing

 Equilibrium

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 INTRODUCTION
General Senses Special Senses
 Somatic sensations  Smell, taste, vision,
(tactile, thermal, pain,
hearing & equilibrium.
proprioceptive) &
visceral sensations.
 Concentrated in specific
locations in the head.
 Scattered throughout the
body.
 Anatomically distinct
structures.
 Simple structures.
 Complex neural pathway.

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 THE CHEMICAL SENSES
 Taste & Smell

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 Chemical senses…
 Taste & smell allow us to separate pleasant &
nutritious foods from undesirable (lethal) foods.
 Sense of smell allows animals to recognize
proximity of other animals or even individuals
among animals.
 Impulses for smell & taste propagate to limbic
system (higher cortical areas),
 certain odors & tastes can evoke strong
emotional responses or a flood of memories.
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 OLFACTION: SENSE OF SMELL
 Gustation (taste) & olfaction (smell), their
chemoreceptors respond to chemicals in aqueous
solution
 Taste – to substances dissolved in saliva
 Smell – to substances dissolved in fluids of
nasal membranes
 Olfactory epithelium = organ of smell.
 Olfactory receptors are surrounded & cushioned
by supporting cells.
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 Anatomy of Olfactory Receptors

 It is estimated that humans can recognize about

10,000 different odors.
 nose contains 10-100 million olfactory receptors
for olfaction within olfactory epithelium.
 Olfactory epithelium occupies superior part of
nasal cavity & consists of 3 kinds of cells:
l Olfactory receptors
l Supporting cells &
l Basal cells.
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 Structure of the Olfactory System

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 Olfactory receptors
 1st order bipolar neurons of olfactory pathway.
 Olfactory hairs (cilia), parts of olfactory receptors
that respond to inhaled chemicals (ODORANT)
• chemicals with odor that can stimulate olfactory
hairs by producing generator potential
(initiating olfactory response).

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Olfactory Transduction

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 Supporting cells
 Provide physical support, nourishment & electrical
insulation for olfactory receptors & detoxify chemicals
that come in contact with the olfactory epithelium.
 Basal cells
 stem cells located b/n bases of supporting cells.
 continually undergo cell division to produce new
olfactory receptors which live for only a month or
so before being replaced.

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 Olfactory (Bowman’s) glands
 Produce mucus to moistens surface of olfactory
epithelium & dissolves odorants so that transduction
can occur.
 Supporting cells & olfactory glands are innervated
by autonomic neurons of facial nerve,
 when stimulated by certain chemicals impulses in
these nerves in turn stimulate lacrimal glands in
eyes & nasal mucous glands.
 result in tears & runny nose after inhaling substances
such as pepper or vapors of household ammonia.
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 Physiology of Olfaction
 Olfactory receptors react to odorant molecules,
generator potential (depolarization) develops &
triggers one or more nerve impulses.
 odorant binds to olfactory receptor protein in plasma
membrane of olfactory hair.
 Olfactory receptor protein is coupled to membrane
protein (G protein), which activates enzyme
adenylate cyclase:
 production of cAMP → opening of Na channels →
inflow of Na → depolarizing GP → generation of nerve
impulse & propagation along axon of olfactory
receptor.
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The Olfactory Pathway

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 On each side of nose 40 bundles of axons of
olfactory receptors form R & L olfactory nerves &
terminate in olfactory bulbs.
 axon terminals of olfactory receptors form synapses with
olfactory bulb neurons in olfactory pathway.
 Axons of olfactory bulb neurons form olfactory tract.
 Some axons of olfactory tract project to 1ry olfactory area
of CC(temporal lobe),conscious awareness of smell.
 From 1ry olfactory area, pathways extend to orbitofrontal
area/ area 11, odor identification & discrimination.
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 Olfactory sensations are only sensations that reach
cerebral cortex without synapsing thalamus.
 Other axons of olfactory tract project to limbic
system & hypothalamus (emotional & memory
evoked responses to odors).
 sexual excitement on smelling certain perfume,
 nausea on smelling food that once made you violently
ill,
 odor evoked memory of a childhood experience.

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 Olfactory Bulb
 Organ which houses all nerves which receive inputs from
olfactory receptors.

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 Olfactory mucosa is directly linked to olfactory bulb
which sends information to frontal lobe for identification
 Collaterals (copies of signals) are send to 3 other
structures
 Amygdala - where emotions are processed
 Hippocampus - where memories are processed
 Hypothalamus - processing involved in eating, drinking &
sexual behavior.
 Smell is strongly associated with emotions, memory,
eating, drinking & sexual behavior.

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 Very Old, Less Old & Newer Olfactory
Pathways into CNS

 Medial olfactory
area represents
very old
olfactory system.
 Lateral olfactory
area is input to
less old &
newer system.

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 Very old olfactory system
 subserve basic olfactory reflexes (lips licking, salivation &
other feeding responses caused by smell of food or by
primitive emotional drives associated with smell).
 Less old system
 provides automatic but partially learned control of food
intake & aversion to toxic & unhealthy foods.
 Newer system
 comparable to most of other cortical sensory systems & is
used for conscious perception & analysis of olfaction.
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 In addition to basic chemical mechanism by which
olfactory cells are stimulated, several physical factors
affect degree of stimulation.
 only volatile substances that can be sniffed into nostrils
can be smelled.
 stimulating substance must be at least slightly water
soluble so that it can pass through mucus to reach
olfactory cilia.
 it is helpful for substance to be at least slightly lipid
soluble, presumably b/c lipid constituents of cilium itself
are weak barrier to non-lipid-soluble odorants.

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 Odor Thresholds
 Olfaction has low threshold, only few molecules of
certain substances need be present in air to be
perceived as an odor.
 Methyl mercaptan (smells like rotten cabbage) can
be detected in concentrations as low as 1/25 billionth
of a mg/ml of air.
 natural gas used for cooking & heating is odorless but
lethal & potentially explosive if it accumulates,
 small amount of methyl mercaptan is added to natural gas
to provide olfactory warning of gas leaks from a pipeline.

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 Odor Adaptation
 Decreasing sensitivity to odors occurs rapidly.
 Olfactory receptors adapt by 50% in 1st sec or so after
stimulation but adapt very slowly thereafter.
 complete insensitivity to certain strong odors occurs about
a minute after exposure.
 Decrease sensitivity involves adaptation process in CNS.
 Large numbers of centrifugal nerve fibers pass from
olfactory regions of brain backward along olfactory tract &
terminate on special inhibitory cells in olfactory bulb
(granule cells).
 after onset of olfactory stimulus, CNS quickly develops
strong feedback inhibition to suppress relay of smell signals
through olfactory bulb.
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 Amoore’s 7 basic smells
1. Camphoraceous
2. Pungent
3. Floral
4. Ethereal
5. Minty
6. Musky
7. Putrid
 all other smells would be a combination of these,
later Amoore had 14 basics, then 21 basics, etc..
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 Henning’s Odor Prism
a graph of 6 basic smells and their possible combinations
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 Functions of Olfaction
 Many animals are macrosmatic
 having a keen sense of smell that is necessary for survival
 Humans are microsmatic
 a less keen sense of smell that is not crucial to survive
 Rats are 8-50 times more sensitive to odors than humans
 Dogs are 300-10,000 times more sensitive
 The difference lies in the number of receptors they each have
 Humans have 10 million &
 Dogs have 1 billion olfactory receptors

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 “Affective Nature of Smell”
(of emotional expression)
 Smell, even more so than taste, has affective quality of
either pleasantness/unpleasantness.
 Smell is probably even more important than taste for the
selection of food.
 person who has previously eaten food that disagreed with
him/her is often nauseated by smell of that same food on
a second occasion.
 perfume of right quality can wreak havoc with human
emotions.
 In some lower animals, odors are primary excitant of
sexual drive.

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 Pheromones: Mammals
 Chemical secreted by animals having;
 Powerful effects on behavior, specifically sexual
behavior, territorial behavior & identification of kin.
Mammals
 Most mammals only become sexually aroused in the
presence of pheromones
 Increased likelihood of pregnancy
 Synchronization of estrus cycles
 Mutual recognition of mother and offspring
 Territory marking (e.g. dogs)

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 Pheromones: Humans
 Infants can correctly identify their own mother's milk & are
much more likely to nurse when its their own mother.
 Female menstrual cycles can be altered by pheromones - the
sorority effect.
 Increases male perception of female attractiveness.
 Increases willingness of females to initiate social contact with
males
 females exposed to alpha androstenol were much more
receptive to male-initiated contact
 more likely to seek out male company than female.
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 GUSTATION: SENSE OF TASTE
 5 primary tastes can be distinguished;
 sour, sweet, bitter, salty and umami.
 Umami (meaty/savory), more recently discovered
tastes by Japanese scientists
 arise from taste receptors that are stimulated by
monosodium glutamate (MSG), substance naturally
present in many foods & added to others as a flavor
enhancer.
 All other flavors (chocolate, pepper & coffee), are
combinations of 5 primary tastes, plus
accompanying olfactory & tactile (touch) sensations.
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 Taste is mainly a function of taste buds in mouth, but it
is common experience that one’s sense of smell also
contributes strongly to taste perception.
 texture of food (detected by tactual senses of mouth) &
presence of substances in food that stimulate pain
endings (pepper) greatly alter taste experience.
 taste is 80% smell
 Importance of taste lies in fact that it allows a person
 to select food in accord with desires & often in accord with
body tissues’ metabolic need for specific substances.

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 Odors from food can pass upward from mouth
into nasal cavity & stimulate olfactory receptors.
 Olfaction is much more sensitive than taste,
 a given concentration of food substance may stimulate
olfactory system 100s x more strongly than it stimulates
gustatory system.
 when you have cold/suffering from allergies & cannot taste
your food,
food it is actually olfaction that is blocked, not taste.

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 Anatomy of Taste Buds & Papillae

 Taste receptors are located in taste buds most

of them are on tongue, but some are found on
 soft palate, pharynx (throat) & epiglottis.
 Each taste bud is oval body consisting of 3
kinds of epithelial cells:
l Supporting cells,
l Gustatory receptor cells &
l Basal cells.
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 Supporting cells
 surround about 50 gustatory receptor cells in
each taste bud.
 Basal cells
 stem cells at periphery of taste bud near
connective tissue layer.
 produce supporting cells, which then develop into
gustatory receptor cells.

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 Gustatory receptor cells
 Has life span of 10 days,
 it doesn’t take taste sensors on tongue too long to recover
from being burned by too hot cup of coffee/cocoa
 Gustatory hair projects to external surface through
taste pore (taste bud opening).
 Synapse with dendrites of 1st order neurons that form
1st part of gustatory pathway,
 dendrites of each 1st order neuron branch profusely &
contact many gustatory receptor cells in several taste
buds.
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 Location of Taste Buds
 Taste buds are found in papillae (elevations on tongue), which
 SA & provide rough texture to upper surface of tongue.
1. Vallate papillae (wall like)
 inverted V-shaped row at back of tongue, houses 100-300
taste buds.
2. Fungiform papillae (mushroom shaped)
 scattered over entire surface of tongue that contain about 5
taste buds each.
3. Foliate papillae (leaf like)
 located in lateral margins of tongue, but most of their taste
buds degenerate in early childhood.

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 Entire surface of tongue has filiform papillae.
 threadlike contain tactile receptors but no taste buds.
  friction b/n tongue & food, making it easier for tongue to
move food in oral cavity.
 Additional taste buds are located on palate, tonsillar
pillars, epiglottis & in proximal esophagus.
 Adults have 3,000-10,000 taste buds & children have few
more.
 Beyond age 45, many taste buds degenerate, causing taste
sensation to become progressively less critical in old age.
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 Taste buds = organs of taste
Functional structures
• Papillae - small rounded ones at the front and sides
- large ones to the back
- Each papilla has one - several hundred
microscopic taste buds

• Taste bud, each has - 50 – 150 taste receptor cells or taste cells
- a person has 2000 – 5000 taste buds

• Taste cells - are only 1% of the tongue epithelium

- taste cells have basal cells that surround the
taste cells and a set of gustatory afferent axons

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Taste Bud Close Up

Molecules of food stimulate the taste cells to send messages to brain

 sweet and salty buds - least sensitive
 bitter ones - most sensitive
Structure of the Taste System

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 Tongue Localization

Its not this simple

Sensitivity differs in different areas,

but all tastes can be perceived at most
areas of the tongue

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 Physiology of Gustation
 TASTANTS = chemicals that stimulate gustatory
receptors.
 once tastant is dissolved in saliva it can make contact
with plasma membrane of gustatory hairs (sites of
taste transduction).
 Receptor potential stimulates exocytosis of
synaptic vesicles from the gustatory receptor cell.
 liberated NT molecules trigger nerve impulses in
1st order sensory neurons that synapse with gustatory
receptor cells.
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 Dissolution in Saliva

Attachment to
Receptors

Generator Potential

Action Potential

THE 4 “BASIC” TASTES ARE SALTY, SOUR, SWEET AND BITTER,

Also: UMAMI (MSG)?, METALLIC? FAT? AMINO ACID?

Different ions, different receptors for different tastes

 Receptor potential arises differently for different
tastants.
 Na+ in salty food enter gustatory receptor cells via
Na channels in the plasma membrane.
 accumulation of Na inside causes depolarization that
leads to release of NT.
 H+ in sour tastants flow into gustatory receptor cells
via H channels.
 opening & closing of other types of ion channels &
result in depolarization & liberation of NT.
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 Tastants stimulating sweet, bitter & umami tastes,
do not themselves enter gustatory receptor cells.
 bind to receptors on plasma membrane that are
linked to G proteins.
 G proteins then activate several different chemicals
known as 2nd messengers inside gustatory receptor
cell.
 different 2nd messengers cause depolarization in
different ways, but result is the same - release of NT.

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 If all tastants cause release of neurotransmitter from
many gustatory receptor cells, why do foods taste
different?
 patterns of nerve impulses in groups of 1st order taste
neurons that synapse with gustatory receptor cells.
 Different tastes arise from activation of different
groups of taste neurons.
 although each individual gustatory receptor cell responds
to > 1 of 5 primary tastes, it may respond more strongly
to some tastants than to others.
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Directly
pass
through
ion
channels
salt &
sour

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 Bind to
membrane
receptors
that activate
second
messenger
systems
that, in turn,
open or close
ion channels
sweet,
umami &
bitter

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 The Gustatory Pathway
 3 cranial nerves contain axons of 1st order
gustatory neurons that innervate taste buds.
 Facial ,,, anterior 2/3 of tongue.
 Glossopharyngeal ,,, posterior 1/3 of tongue.
 Vagus ,,, throat & epiglottis.

 From taste buds, nerve impulses propagate

along these cranial nerves to gustatory nucleus
in medulla oblongata.

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 From medulla;
 some axons carrying taste signals project to
limbic system & hypothalamus and
 others project to thalamus.

 Taste signals that project from thalamus to 1ry

gustatory area in parietal lobe of cerebral
cortex (area 43)
 give rise to conscious perception of taste.

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 Taste Reflexes Integrated in Brain Stem
 From the tractus solitarius,
 many taste signals are transmitted within brain
stem itself directly into superior & inferior
salivatory nuclei, and
• these areas transmit signals to submandibular,
sublingual & parotid glands
 to help control the secretion of saliva
during the ingestion & digestion of food.

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 Taste Thresholds & Adaptation
 Threshold for taste varies for each of 1 ry tastes.
 Threshold for bitter substances (quinine) is lowest.
 Poisonous substances often are bitter, low threshold
(high sensitivity) have protective function.
many deadly toxins found in poisonous plants are
alkaloids & virtually all of these cause intensely bitter
taste, usually followed by rejection of the food.
 Threshold for sour substances (lemon) as measured
by using HCl is somewhat higher.

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 Thresholds for salty substances (NaCl) & sweet
substances (sucrose) are similar & are higher
than those for bitter or sour substances.
 Complete adaptation to a specific taste can
occur in 1–5 minutes of continuous stimulation.
 Taste adaptation is due to changes that occur in
 taste & olfactory receptors and
 neurons of gustatory pathway in CNS.

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 Taste Preference & Control of Diet

 Taste preference
 animals choose certain types of food in
preference to others.
 animal
automatically uses this to help control
type of diet it eats.
 Taste preferences often change in accord with
the body’s need for certain specific substances.

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 Adrenalectomized (salt depleted animals)
 automatically select drinking water with high concentration
of NaCl in preference to pure water & this is often sufficient
to supply needs of body & prevent salt-depletion death.
 Insulin injecteted (blood sugar depleted animals)
 automatically chooses sweetest food from among many
samples.
 Parathyroidectomized (Ca++ depleted animals)
 automatically choose drinking water with high concentration of
calcium chloride.
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 Same phenomena are also observed in everyday
life, for instance,
 “salt licks” of desert regions are known to attract
animals from far and wide.
 Human beings reject any food that has an
unpleasant affective sensation,
 which in many instances protects our bodies
from undesirable substances.

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Survival Significance of Taste

Taste Stimulus Significance

Complex organic
Sweet molecules energy
(sugars and carbohydrates)

Salt NACL Nerve conduction

Spoiled food
Sour Acids (H+) or vitamins
Organic alkaloids
Bitter Metallic salts poison

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 VISION
 Act of seeing, extremely important to human survival
 More than half sensory receptors in human body are
located in eyes & large part of cerebral cortex is
devoted to processing visual information.
 In this section we examine
 electromagnetic radiation
 accessory structures of eye, eyeball
 formation of visual images, physiology of vision &
visual pathway from eye to brain.
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 Electromagnetic Radiation
 Energy in form of waves that radiates from sun.
 Types of EMR; γ rays, x-rays, UV rays, visible
light, infrared radiation, micro & radio waves.
 Rnge of EMR = electromagnetic spectrum.
 Wavelength, distance b/n 2 consecutive peaks of
EMW.
 range from short (gamma rays) to long (radio
waves) ; smaller than nanometer & greater than
meter.
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 Eyes are responsible for detection of visible light,
400-700 nm.
 Visible light exhibits colors that depends on wave length
(violet 400 nm & 700 nm red).
 Object can absorb certain wl of visible light & reflect
others; object will appear color of wl that is reflected.
 green apple appears green b/c it reflects mostly green light
& absorbs most other wl of visible light.
 white object reflects all wl of visible light.
 black object absorbs all wl of visible light.

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 Accessory Structures of Eye
 Eyelashes & Eyebrows
 Protect eyeballs from foreign objects, perspiration & direct
rays of the sun.
 Eyelids/palpebrae
 Shade eyes during sleep, protect eyes from excessive light
& foreign objects.
 Lacrimal caruncle
 Contains sebaceous (oil) glands & sudoriferous (sweat)
glands.
 Meibomian glands
 Secrete fluid that keep eyelids from adhering to each other.
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 The Lacrimal Apparatus

 Group of
structures that
produces &
drains lacrimal
fluid/tears.
 Parasympathetic
fibers of facial
(VII) nerves
supply lacrimal
glands.

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 The fluid is watery solution containing salts, mucus &
lysozyme, protective bactericidal enzyme.
 The fluid protects, cleans, lubricates & moistens eyeball.
 Lacrimation is protective mechanism, as tears dilute
& wash away irritating substance.
 Watery eyes occur when inflammation of nasal
mucosa (cold) obstructs nasolacrimal ducts & blocks
drainage of tears.

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 Extrinsic Eye Muscles
 Eyes sit in bony depressions of skull called orbits.
 protect eyes, stabilize in 3D space & anchor to
muscles that produce their essential movements.
 Are capable of moving eye in almost any direction.
 Superior & inferior rectus, lateral & medial
rectus, superior & inferior oblique.
 They are supplied by cranial nerves III, IV & VI.
 In general, the motor units in these muscles are
small.

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 I. Fibrous Tunic
 Outer layer of eyeball with anterior cornea &
posterior sclera.
 Cornea is transparent coat that covers colored iris,
its curvature helps focus light onto the retina.
 Sclera (hard) “white” of the eye, covers entire eyeball
except cornea.
 gives shape to eyeball, makes it more rigid, protects its
inner parts & site of attachment for extrinsic eye muscles.
 Scleral venous sinus (canal of Schlemm), opening
that drains aqueous humor.
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 II. Vascular Tunic (uvea)
 Middle layer of eyeball, composed of 3 parts:
 choroid, ciliary body & iris .
 Choroid, highly vascularized posterior portion of
vascular tunic, lines most of internal surface of sclera.
 blood vessels provide nutrients to retina.
 dark brown color due to melanocytes (melanin).
 Melanin absorbs stray light rays, prevents reflection
& scattering of light within the eyeball.
 Image cast on retina by cornea & lens remains sharp &
clear.
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 Ciliary body
 Anterior portion of vascular tunic, consists of ciliary
processes & ciliary muscle.
 Ciliary processes are protrusions or folds on internal
surface of ciliary body that contain
 blood capillaries that secrete aqueous humor &
 zonular fibers (suspensory ligaments) attach to lens.
 Ciliary muscle is circular band of smooth muscle.
 contraction/relaxation of ciliary muscle changes the
tightness of zonular fibers, which alters shape of lens,
adapting it for near or far vision.

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 Iris (rainbow)
 Colored portion of eyeball, flattened donut shape.
 Suspended b/n cornea & lens & is attached at its outer
margin to ciliary processes.
 Consists of melanocytes & circular & radial smooth
muscle fibers.
 Amount of melanin in iris determines eye color.
 eyes appear brown to black when iris contains large
amount of melanin, blue when melanin is very low & green
when melanin is moderate.
 Principal function is to regulate amount of light entering
eyeball through pupil.

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 Pupil(little person)
 Hole in center of iris.
 Appears black b/c, as you look through lens, you see
heavily pigmented back of the eye (choroid & retina).
 If bright light is directed into pupil, reflected light is
red b/c of blood vessels on surface of retina.
 a person’s eyes appear red in photograph (“red eye”)
when flash is directed into the pupil.
 Autonomic reflexes regulate pupil diameter in
response to light levels.

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 III. Retina
 Inner layer of eyeball, lines posterior 3/4 of eyeball &
beginning of visual pathway.
 Can be viewed with ophthalmoscope,
 instrument that shines light into eye & allows observer
to peer through pupil providing magnified image of
retina & its blood vessels & optic (II) nerve.
 Surface of retina is only place in body where blood
vessels can be viewed directly & examined for
 pathological changes (hypertension, DM, cataracts & age-
related macular disease).
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 Optic disc (blind spot) is site where optic (II) nerve exits
the eyeball.
 Central retinal artery fan out to nourish anterior surface
of retina; central retinal vein drains blood from retina
through optic disc.
 Retina consists of pigmented & neural layer.
 Pigmented layer is melanin-containing epithelial cells
located b/n choroid & neural part of retina.
 Neural (sensory) layer is multilayered outgrowth of brain
that processes visual data extensively before sending
nerve impulses into axons that form optic nerve.

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 Layers of retinal neurons
 Photoreceptor, bipolar & ganglion cell.
 separated by outer & inner synaptic layers where synaptic
contacts are made.
 Light passes through ganglion & bipolar cell layers &
both synaptic layers before it reaches photoreceptor.
 Horizontal & amacrine, types of cells present in
bipolar cell layer of the retina.
 form laterally directed neural circuits that modify signals
being transmitted along the pathway from photoreceptors to
bipolar cells to ganglion cells.
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 Photoreceptors
 Specialized cells that begin process by which light rays
are ultimately converted to nerve impulses.
 Two types of photoreceptors: rods & cones.
 Each retina has about 6 million cones & 120 million rods.
 Rods allow us to see in dim light, such as moonlight.
 B/c rods do not provide color vision, in dim light we can see
only black, white & gray.
 Cones stimulated by brighter lights which produce color
vision.

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 Types of cones

 3 types of cones are present in the retina:

 blue cones, which are sensitive to blue light,
 green cones, which are sensitive to green light, and
 red cones, which are sensitive to red light.
 Color vision results from stimulation of various
combinations of these 3 types of cones.
 Most of our experiences are mediated by cone system,
loss of which produces legal blindness.
 Person who loses rod vision mainly has difficulty
seeing in dim light, should not drive at night.

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 Lens
 Located behind pupil & iris, within eyeball cavity.
 Crystallins proteins arranged like layers of onion,
 make up refractive media of lens, which normally is
perfectly transparent & lacks blood vessels.
 Enclosed by clear connective tissue capsule & held
in position by encircling zonular fibers, which attach
to ciliary processes.
 Helps focus images on the retina to facilitate clear
vision.

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 Interior of the Eyeball
 Aqueous humor
 transparent watery fluid that nourishes lens & cornea,
replaced every 90min.
 Vitreous body
 transparent, giving retina even surface for reception of clear
images.
 does not undergo constant replacement, formed during
embryonic life & consists of mostly water plus collagen
fibers & hyaluronic acid.
 contains phagocytic cells that remove debris, keeping this
part of eye clear for unobstructed vision.
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 Intraocular pressure (16 mmHg), is produced
mainly by aqueous humor & partly by vitreous body.
 maintains shape of eyeball & prevents it from
collapsing.
 Puncture wounds to the eyeball may cause the loss
of aqueous humor and the vitreous body.
 This in turn causes a decrease in intraocular
pressure, a detached retina & in some cases
blindness.

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 Image Formation
 In some ways eye is like camera:
 Its optical elements focus image of some object on light-
sensitive “film”—retina—while ensuring correct amount of
light to make proper “exposure.”
 To understand how eye forms clear images of
objects on retina, we must examine 3 processes:
¢ refraction, bending of light by lens & cornea;
¢ accommodation, change in shape of lens
¢ constriction, narrowing of the pupil.

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 Refraction of Light Rays
 As light rays enter the eye, they are refracted at
anterior & posterior surfaces of cornea.
 Both surfaces of lens further refract light rays so they come
into exact focus on retina.
 Images focused on retina are
 inverted (upside down), reversal (right-to-left); light from
right side of object strikes left side of retina & vice versa.
 The reason world does not look inverted & reversed
is that brain “learns” early in life to coordinate visual
images with orientations of objects.
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 75% of refraction of light occurs at cornea.
 Lens provides 25% of focusing power & changes focus to
view near/distant objects.
 Objects ≥ 6m away from viewer, light rays reflected are
nearly parallel to one another.
 lens must bend these parallel rays just enough so that they
fall exactly focused on central fovea (sharpest vision).
 Objects < 6m are divergent rather than parallel, rays must
be refracted more if they are to be focused on retina.
 this additional refraction is accomplished through a process
called accommodation.
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 Accommodation &
Near Point of Vision
 Concave, curves inward, like inside of hollow ball.
 causes light rays to refract away from each other.
 Convex, curves outward like surface of ball.
 refract incoming light rays toward each other, so that they
eventually intersect.
 lens of eye is convex on its both surfaces, its focusing
power ↑ as its curvature becomes greater.
 when eye is focusing on close object, lens becomes more
curved, causing greater refraction of light rays.
↑in curvature of lens for near vision = accommodation.
 Near point of vision (young adult 10cm), minimum distance
from eye that object can be clearly focused with maximum
accommodation.
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 How does accommodation occur?

 When you are viewing distant objects,

 ciliary muscle relax & lens is flatter b/c it is stretched in all
directions by taut zonular fibers.
 When you view a close object,
 ciliary muscle contracts, zonular fibers relax & lens
becomes more spherical (more convex), ↑its focusing
power & causes greater convergence of light rays.
 PNS of oculomotor (III) nerve innervate ciliary
muscle & mediate process of accommodation.
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 Refraction Abnormalities
 Emmetropic eye (normal eye)
 refract light rays from object 6m away so that clear image
is focused on retina.
 Myopia (nearsightedness)
 too long eyeball/ thicker lens, image in front of retina.
 can see close objects clearly, but not distant objects.
 Hyperopia (farsightedness/hypermetropia)
 Short eyeball/thinner lens, image behind retina.
 can see distant objects clearly, but not close ones.
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 Physiology
of Vision
Photoreceptors
• Rods &
• Cones
Photopigments
• Colored proteins in
plasma membrane of
outer segment of
photoreceptors.

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 Photopigments
 Undergoes structural changes when it absorbs light.
 Light absorption initiates events that lead to production of
receptor potential.
 Rhodopsin, single type of photopigment in rods.
 Cone photopigments , present in retina, one in each of
the 3 types of cones (BGR).
 Color vision results from different colors of light selectively
activating different cone photopigments.
 Contain OPSIN (glycoprotein) & RETINAL (vita. A derivative)

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 Retinal, light absorbing part of visual photopigments.
 Vita. A derivatives are formed from carotene, plant
pigment that gives carrots their orange color.
 Good vision depends on adequate dietary intake of
carotene-rich vegetables (carrots, spinach, broccoli &
yellow squash) or foods that contain vita.A, such as liver.
 Opsins, 4 types, 3 in cones & 1 in rods (rhodopsin).
 Small variations in aa sequences of different opsins permit
rods & cones to absorb different colors of incoming light.

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 Rod disc in outer segment

Rhodopsin
molecule Light
Light
Disc cis-
membrane retinal
4 Cis-retinal
binds to opsin opsin 1 Isomerization
(regeneration) of retinal
Colored
cis- photopigment trans-
retinal (rhodopsin) retinal

trans-
opsin retinal opsin

3 Retinal 2 Trans-retinal
isomerase separates
converts trans- from opsin
to cis-retinal opsin (bleaching)

Colorless products
 Pigmented layer of retina adjacent to photoreceptors
stores large quantity of vita.A & contributes to
regeneration process in rods.
 Extent of rhodopsin regeneration decreases drastically if
retina detaches from the pigmented layer.
 Cone photopigments regenerate much more quickly than
rhodopsin & are less dependent on pigmented layer.
 After complete bleaching, regeneration of half of rhodopsin
takes 5min; half of cone photopigments regenerate in only
90sec.
 Full regeneration of bleached rhodopsin takes 30-40min.
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 Release of NT by Photoreceptors

 Absorption of light & isomerization of retinal initiates

chemical changes in photoreceptor outer segment
that lead to production of receptor potential.
 In darkness, Na flow into photoreceptor outer
segments through L-gated Na channels/cGMP.
 Inflow of Na, “dark current,” partially depolarizes the
photoreceptor.
 in darkness membrane potential of photoreceptor is 30
mV, much closer to 0 than typical neuron’s RMP of 70 mV.

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 Partial depolarization during darkness triggers
continual release of NT at synaptic terminals.
 NT in rods & perhaps in cones, glutamate (glutamic acid).
 At synapses b/n rods & some bipolar cells, glutamate
is inhibitory NT:
 triggers IPSPs that hyperpolarize bipolar cells & prevent
them from sending signals to ganglion cells.
 Light strikes retina & cis-retinal undergoes isomerization,
enzymes are activated that break down cGMP.

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 Some cGMP-gated Na channels close, ↓Na inflow, &
membrane potential becomes more -ve,(70mV).
 produces hyperpolarizing receptor potential that
decreases release of glutamate.
 Dim lights cause small & brief receptor potentials that
partially turn off glutamate release; brighter lights elicit
larger & longer receptor potentials that more completely
shut down NT release.
 light excites bipolar cells that synapse with rods by turning
off release of inhibitory NT.
 Excited bipolar cells subsequently stimulate ganglion
cells to form action potentials in their axons.
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 The Visual Pathway
 Visual signals in retina undergo considerable
processing at synapses among various types of
neurons in retina
 Horizontal cells
 Bipolar cells
 Amacrine cells

 Axons of retinal ganglion cells provide output from

retina to brain, exiting eyeball as the optic (II)
nerve.
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 Processing of Visual Input
in Retina
 Once receptor potentials arise in rods & cones, they
spread through inner segments to synaptic terminals.
 NT molecules released by rods & cones induce local GPs
in both bipolar & horizontal cells.
 6- 600 rods synapse with 1 bipolar cell; cone more often
synapses with 1 bipolar cell.
 Convergence of many rods onto 1 bipolar cell increases
light sensitivity of rod vision but slightly blurs image.
 Cone vision, although less sensitive, is sharper because
of 1-1 synapses b/n cones & their bipolar cells.

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Horizontal cells Amacrine cells
 Transmit inhibitory signals to  Excited by bipolar cells,
bipolar cells in areas lateral synapse with ganglion cells
to excited rods & cones. &
 Lateral inhibition enhances  transmit information to
contrasts in visual scene b/n signal change in level of
areas of retina that are illumination of retina.
strongly stimulated & weakly  Bipolar/amacrine cells
stimulated. transmit excitatory signals to
 assist in differentiation of ganglion cells to depolarize
various colors. & initiate nerve impulses.

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 Brain Pathway & Visual Fields
 Optic (II) nerve pass through optic chiasm.
 Some axons cross to opposite side but others remain
uncrossed.
 Optic tract enter brain & most of them terminate in
LGN of thalamus.
 synapse with neurons whose axons form optic radiations,
which project to 1ry visual areas in occipital lobes of
cerebral cortex (area 17) & visual perception begins.
 Some of fibers in optic tracts terminate in
 superior colliculi, control extrinsic eye muscles &
 pretectal nuclei, pupillary & accommodation reflexes.
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 Visual field= everything that can be seen by 1 eye.
 our eyes are located anteriorly in our heads, visual fields
overlap (binocular visual field).
 Visual field of each eye is divided into 2 regions:
nasal/central & temporal/peripheral half.
 For each eye,
 light rays from object in nasal half of visual field fall on
temporal half of retina &
 light rays from object in temporal half of visual field fall on
nasal half of retina.

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 Visual information from right half of each visual field is
conveyed to left side of brain & visual information from
left half of each visual field is conveyed to right side of
brain, as follows :
1. Axons of all retinal ganglion cells in 1 eye exit eyeball at
optic disc & form optic nerve on that side.
2. At optic chiasm, axons from temporal half of each retina
do not cross but continue directly to LGN of thalamus on
same side.
3. Axons from nasal half of each retina cross optic chiasm &
continue to opposite thalamus.

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 Visual field of Visual field of
left eye right eye

Temporal Nasal Nasal Temporal

half half half half

Left eye Right eye

Nasal retina
Temporal Nasal Temporal
retina 1 retina retina
3 1
3
4 Optic 4
2 tract 2
Midbrain Midbrain
5 5
Lateral geniculate nucleus
of the thalamus
Optic 6 Optic
radiations 6
radiations

Primary visual area of cerebral

cortex (area 17) in occipital lobe

Left eye and its pathways Right eye and its pathways
4. Each optic tract (crossed & uncrossed axons)
project from optic chiasm to thalamus on 1 side.
5. Axon collaterals of retinal ganglion cells project to
midbrain, govern constriction of pupils in response
to light & coordination of head & eye movements.
6. Collaterals also extend to suprachiasmatic nucleus
of hypothalamus,
4. establishes patterns of sleep & other activities that occur
on circadian/daily schedule in response to intervals of light
& darkness.

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6. Axons of thalamic neurons form optic radiations as
they project from thalamus to 1ry visual area of
cortex on same side.
7. Although we have just described visual pathway as
single pathway, visual signals are thought to be
processed by at least 3 separate systems in
cerebral cortex, each with its own function.
6. shape of objects,
7. regarding color of objects &
8. about movement, location & spatial organization.

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 HEARING & EQUILIBRIUM
 Hearing = the ability to perceive sounds.
 Ear is an engineering marvel, its sensory receptors
can transduce sound vibrations with amplitudes as
small as diameter of atom of gold (0.3nm) into
 electrical signals 1000 times faster than photoreceptors
can respond to light.
 Ear also contains receptors for equilibrium,
 sense that helps you maintain your balance & be aware of
your orientation in space.

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 Anatomy of the Ear

 Ear is divided into 3 main regions:

1. External ear
 collects sound waves & channels them inward

2. Middle ear
 conveys sound vibrations to the oval window
¢ Internal ear
 houses the receptors for hearing & equilibrium.

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 External (Outer) Ear
 Auricle/pinna, external auditory canal & tympanic
membrane/eardrum.
 Perforated eardrum
 tearing of tympanic membrane may be due to
pressure from cotton swab, trauma, or middle ear
infection & usually heals within a month.
 Otoscope
 viewing instrument that illuminates & magnifies
external auditory canal & tympanic membrane.
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 Combination of hairs & cerumen helps prevent dust
& foreign objects from entering the ear.
 Cerumen also prevents damage to delicate skin of
external ear canal by water & insects.
 usually dries up & falls out of the ear canal.
 Some people produce large amount of cerumen,
which can become impacted & can muffle incoming
sounds.
 treatment for impacted cerumen is usually periodic ear
irrigation/removal of wax with blunt instrument by trained
medical personnel.

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 Middle Ear
 Small, air-filled cavity in petrous portion of
temporal bone.
 Separated from external ear by tympanic
membrane & from internal ear by oval & round
window.
 Auditory ossicles (smallest bones in the body )
are connected by synovial joints.
 malleus, incus & stapes, commonly called hammer, anvil &
stirrup, respectively.
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 Tensor tympani muscle,
 supplied by trigeminal (V) nerve, limits movement & ↑
tension on eardrum to prevent damage to inner ear from
loud noises.
 Stapedius muscle (smallest skeletal muscle in human)
 supplied by facial (VII) nerve, by dampening large
vibrations of stapes due to loud noises it protects oval
window, but it also ↓ sensitivity of hearing.
 B/c it takes fraction of second for tensor tympani &
stapedius muscles to contract,
 they can protect inner ear from prolonged loud noises but
not from brief ones such as a gunshot.

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 Auditory tube
(pharyngotympanic/eustachian tube)
 Connects middle ear with nasopharynx (throat).
 Closed at pharyngeal end, during swallowing & yawning it
opens allowing air to enter/leave middle ear until pressure
in middle ear equals atmospheric pressure.
 When pressures are balanced, tympanic membrane
vibrates freely as sound waves strike it.
 If pressure is not equalized, intense pain, hearing
impairment, ringing in ears & vertigo could develop.
 Route for pathogens to travel from nose & throat to
middle ear, causing most common type of ear
infection (otitis media).
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 Internal (Inner) Ear
 Also called labyrinth b/c of its complicated series of
canals.
 Bony labyrinth divided into 3 areas: semicircular
canals, vestibule & cochlea.
 contains perilymph(CSF) & surrounds membranous
labyrinth.
 Membranous labyrinth, tubes inside bony labyrinth
 contains endolymph & house receptors for hearing &
equilibrium.
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133 05/05/24
 Vestibule = central portion of bony labyrinth.
 Utricle (little bag) & saccule (little sac) are
located in vestibule of membranous labyrinth.
 Semicircular canals, projecting superior &
posterior from vestibule.
 ampulla (saclike duct) swollen enlargement at
one end of each canal.
 Semicircular ducts, portions of membranous
labyrinth that lie inside bony SCCs.
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 Vestibular branch of vestibulocochlear (VIII) nerve
consists of ampullary, utricular & saccular nerves.
 The nerves contain both 1st order sensory & motor
neurons that synapse with receptors for equilibrium.
 sensory neurons carry sensory information from receptors
 motor neurons carry feedback signals to receptors,
apparently to modify their sensitivity.
 Cell bodies of sensory neurons are located in the
vestibular ganglia.
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136 05/05/24
 Cochlea
 Bony spiral canal resembles snail’s shell & makes 3
turns around central bony core modiolus.
 Divided into 3 channels:
 scala vestibuli
 scala media (cochlear duct)
 scala tympani.
 Vestibular membrane separates cochlear duct from
scala vestibule & basilar membrane separates
cochlear duct from scala tympani.
 Resting on basilar membrane, spiral organ/organ of Corti

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Spiral organ
coiled sheet of
epithelial cells,
including:
Supporting
cells
Hair cells
(hearing
receptors), 16,000.

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 Hair cells (hearing receptors)
 Two groups of hair cells:
 inner hair cells arranged in 1 row
 outer hair cells arranged in 3 rows
 At apical tip of each hair cell are 40–80 stereocilia that
extend into endolymph of cochlear duct.
 long, hairlike microvilli arranged in several rows of graded
height.
 Inner & outer hair cells synapse 1st order sensory & motor
neurons from cochlear branch of vestibulocochlear nerve.
 sensory neurons cell bodies are located in spiral ganglion.

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 Outer hair cells outnumber inner hair cells by 3 to 1,
 inner hair cells synapse with 90–95% of 1st order sensory
neurons in cochlear nerve that relay auditory information to
the brain.
 90% of motor neurons in cochlear nerve synapse with outer
hair cells.
 Tectorial membrane, flexible gelatinous membrane,
covers hair cells of spiral organ.
 ends of stereocilia of hair cells are embedded in tectorial
membrane while bodies of hair cells rest on basilar
membrane.
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 Nature of Sound Waves
 Sound waves are alternating high & low-pressure
regions traveling in same direction through some medium
(such as air).
 originate from vibrating object in much same way that
ripples arise & travel over surface of pond when you toss
stone into it.
 Pitch , frequency of sound vibration.
 higher frequency of vibration, higher pitch.
 Sounds heard by human ear, 500 & 5000 hertz, entire
audible range extends from 20- 20,000 Hz.
 Sounds of speech b/n 100 & 3000 Hz, sounds from jet
plane range 20-100 Hz.
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 Louder sound, larger intensity (size/amplitude) of
vibration.
 Sound intensity is measured in decibels (dB), ↑of 1dB
represents 10x ↑ in sound intensity.
 Hearing threshold, average young adult can
distinguish sound from silence, defined as 0 dB at
1000 Hz.
 rustling leaves have 15db, whispered speech 30; normal
conversation 60, vacuum cleaner 75, shouting 80 & nearby
motorcycle 90db.
 Sound becomes uncomfortable to normal ear at 120
dB & painful above 140 dB.
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 Physiology of Hearing
1. Auricle directs sound waves into
external auditory canal.
2. Sound waves strike tympanic membrane
alternating waves of high & low pressure
in air cause TM to vibrate back & forth.
 TM vibrates slowly in response to low-freq.(pitch) &
rapidly in response to high freq. (pitch) sounds.
3. Vibration from TM is transmitted to malleus, incus &
then to stapes.
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 Malleus Incus Stapes vibrating Helicotrema Cochlea
in oval window

Sound waves
Perilymph

3 8 Scala
7 tympani
4
Scala
5 vestibuli
6 Basilar
1 2 9 membrane
External auditory 8
canal Spiral organ
(organ of Corti)
Tectorial membrane
Vestibular membrane
Cochlear duct
Tympanic (contains endolymph)
membrane
Secondary tympanic
membrane vibrating
Middle ear Auditory tube
in round window
4) As stapes moves back & forth vibrates oval window.
 Vibrations at oval window are 20x more vigorous than
tympanic.
5. Movement of stapes at oval window sets up fluid
pressure waves in perilymph of cochlea.
 As oval window bulges inward, it pushes on perilymph of
scala vestibuli.
6. Pressure waves transmit from scala vestibuli to
 scala tympani & to round window, causing it to bulge
outward into middle ear.
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7. Pressure waves travel through perilymph of scala
vestibuli →vestibular membrane→endolymph inside
cochlear duct.
8. Pressure waves in endolymph cause basilar
membrane to vibrate, which moves hair cells of
spiral organ against tectorial membrane.
7. bending of stereocilia generate nerve impulses in 1st
order neurons in cochlear nerve fibers.
9. Sound waves of various frequencies cause certain
regions of basilar membrane to vibrate more
intensely than other regions.
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 Basilar membrane
 Each segment of basilar membrane is “tuned” for
particular pitch.
 Base of cochlea (oval window), narrower & stiffer high freq.
(pitch) sounds induce maximal vibrations.
 Apex of cochlea, wider & more flexible;
flexible low-freq (pitch)
sounds cause maximal vibration of basilar membrane.
 Loudness is determined by intensity of sound waves
 High-intensity sound waves cause larger vibrations of basilar
membrane, which leads to higher frequency of nerve impulses
reaching the brain.
 stimulate larger number of hair cells, transduce mechanical
vibrations into electrical signals.

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 As basilar membrane vibrates, hair bundles at apex
of hair cell bend back & forth & slide against one
another.
 Tip link protein connects tip of each stereocilium to
M-gated ion channel(transduction channel) in its
taller stereocilium neighbor.
 as stereocilia bend in direction of taller stereocilia, tip links
tug on transduction channels & open them.
 channels allow cations (K) in endolymph to enter hair cell
cytosol & produce depolarizing receptor potential.
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 Depolarization quickly spreads along plasma
membrane & opens V-gated Ca++ channels in base of
hair cell.
 inflow of Ca++ triggers exocytosis of NT (glutamate).
 nerve impulses in 1st order sensory neurons that synapse
with base of hair cell increases.
 Bending of stereocilia in opposite direction closes
transduction channels,
 allows hyperpolarization to occur & reduces NT release
from hair cells, decreases frequency of nerve impulses in
sensory neurons.
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 The Auditory Pathway
 Bending of stereocilia causes release of NT
(glutamate), generates nerve impulses in sensory
neurons.
 Cell bodies of sensory neurons are located in spiral
ganglia.
 Nerve impulses pass along axons of these neurons, which
form cochlear branch of vestibulocochlear (VIII) nerve.
 These axons synapse with neurons in cochlear nuclei
in medulla oblongata on same side.
 Some of axons from cochlear nuclei decussate in medulla,
ascend in lateral meniscus tract on opposite side &
terminate in inferior colliculus in midbrain.
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152 05/05/24
 Other axons from cochlear nuclei end in superior
olivary nucleus in pons on each side.
 Slight d/ces in timing of nerve impulses arriving from 2
ears at superior olivary nuclei allow us to locate
source of sound.
 Axons from superior olivary nuclei also ascend in
lateral meniscus tracts on both sides & end in
inferior colliculi.
 From each inferior colliculus, nerve impulses
conveyed to MGN in thalamus & finally to 1ry
auditory area of cerebral cortex in temporal lobe of
cerebrum (areas 41 & 42).
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 Physiology of Equilibrium
 Vestibular apparatus = equilibrium receptor organs.
 saccule, utricle & semicircular ducts
 Two types of equilibrium/balance.
1. Static equilibrium: maintenance of position of body
(head) relative to force of gravity.
 body movements like tilting head & linear acceleration or
deceleration (elevator /car).
2. Dynamic equilibrium: maintenance of body position
(head) in response to sudden movements
1. rotational acceleration or deceleration.
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155 05/05/24
 Otolithic Organs: Saccule & Utricle
 Macula, small, thickened walls of utricle & saccule.
 are receptors for static equilibrium.
 provide sensory information on position of head in
space & essential for maintaining appropriate posture
& balance.
 detect linear acceleration & deceleration, sensations in
elevator/car (speeding up/slowing down).
 consist of hair cells (sensory receptors) & supporting
cells.

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 Hair cells have on their surface 40-80 stereocilia
plus 1 kinocilium.
 stereocilia are connected by tip links.
 hair bundle (stereocilia & kinocilium).

 Otolithic membrane, supporting cells that

secrete thick, gelatinous, glycoprotein layer.
 holds layer of dense calcium carbonate crystals,
otoliths (ear stones).

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 If you tilt your head forward, otolithic membrane
(otoliths) is pulled by gravity.
 slides “downhill” over hair cells in direction of tilt, bending
hair bundles.
 If you are sitting upright in car that suddenly jerks
forward,
 otolithic membrane lags behind head movement, pulls on
hair bundles & makes them bend in other direction.
 Bending of hair bundles in
 one direction stretches tip links & open transduction
channels, producing depolarizing receptor potentials;
 opposite direction closes transduction channels & produces
hyperpolarization.
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 As hair cells depolarize & repolarize, they release
NT at a faster/slower rate.
 hair cells synapse with 1st order sensory neurons in
vestibular branch of vestibulocochlear (VIII) nerve, fire
impulses at slow/rapid pace depending on NT amount.
 Motor neurons also synapse with hair cells &
sensory neurons.
 regulate sensitivity of hair cells & sensory
neurons.
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 Semicircular Ducts
 Function in dynamic equilibrium.
 Lie at right angles to one another in 3 planes:
 2 vertical (anterior & posterior) & 1 horizontal lateral.
 this positioning permits detection of rotational
acceleration or deceleration.
 Crista, small elevation in ampulla (dilated portion of
each duct).
 contains group of hair cells & supporting cells.
 Cupula mass of gelatinous material covering crista.
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 When you move your head, attached SSDs & hair
cells move with it.
 Endolymph within ampulla, however, is not attached & lags
behind.
 As the moving hair cells drag along the stationary
endolymph, hair bundles bend.
 Bending of hair bundles produces receptor potentials.
 In turn, receptor potentials lead to nerve impulses
that pass along vestibular branch of
vestibulocochlear nerve.
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 Equilibrium Pathways
 Bending of hair bundles of hair cells in SSDs, utricle/saccule
causes release of NT (glutamate), generates nerve impulses in
sensory neurons that innervate hair cells.
 Cell bodies of sensory neurons are located in vestibular
ganglia.
 Nerve impulses pass along axons of these neurons, which form
vestibular branch of vestibulocochlear (VIII) nerve.
 Most of these axons synapse with sensory neurons in vestibular
nuclei, major integrating centers for equilibrium, in medulla
oblongata & pons.
 Vestibular nuclei also receive input from eyes & proprioceptors
(neck & limb muscles) that indicate position of head & limbs.
 Remaining axons enter cerebellum through inferior cerebellar
peduncles.
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 Vestibular nuclei integrate information from vestibular, visual &
somatic receptors & then send commands to
¢ Oculomotor (III), trochlear (IV) & abducens (VI) control
coupled movements of eyes with those of head to help maintain
focus on visual field;
¢ Accessory (XI) nerves control head & neck movements to
assist in maintaining equilibrium;
¢ Vestibulospinal tract, conveys impulses down spinal cord to
maintain skeletal muscle tone to maintain equilibrium
¢ Ventral posterior nucleus in thalamus → vestibular area in
parietal lobe of cerebral cortex (area 1, 2 & 3) to provide
conscious awareness of position & movements of head & limbs.

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 I WISH YOU ALL THE BEST S
E
T E
H U
A
A T
N
Z
K
T
Y O
O P
U GOD BLESS YOU & GOD BLESS ETHIOPIA
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