Dr S Voiculescu

BALANCE

Vestibular system
Vestibular apparatus:

utricle, saccule and

semicircular ducts.
Sensory cells of
vestibular apparatus lie
in the ampular cristae
of canals and maculae
in the utricle and
saccule.

 Both types of organs- hair cells- special stereocilia over

the apical surface and one kinocilium:

In the utricle and saccule, the receptor organ is called

macula- stereocillia and kinocilium are embedded in

cuticular plate with otoliths (CaCO3 crystals)
In the semicircular ducts, the receptor organs are called
ampular cristae- hairs are covered by cupulae, composed
of a gelatinous material similar to otolithic membrane
but lacking otoliths.

Macula (utricular and saccular)

Cristae ampullaris (semicircular

canals)

- Utricle and saccule:

Linear acceleration

detection
Head position
(gravitation)

- Semicircular canals
Angular motion

Function of the maculae

detect head position and linear acceleration
otoliths (small calcium carbonate particles) drag on

the stereocilia when the head changes position

when the body is in anatomical position: the patch of
hair cells in the UTRICLE is nearly horizontal, with
the stereocilia oriented vertically
the sensory epithelium is vertical in the SACCULE,
with the stereocilia oriented horizontally

 when the body is in

anatomical position:
the patch of hair cells
in the UTRICLE is
nearly horizontal,
with the stereocilia
oriented vertically
the sensory
epithelium is vertical
in the SACCULE,
with the stereocilia
oriented
horizontally

 K+ TRPA channels + tip- links

Depolarisation- lean towards the kinocillium- K+ ch
open
Hyperpolarisation- lean towards the stereocillia- K+ ch
close

 orientation of the

stereocilia within the

sensory epithelium is
determined by the
STRIOLA, a curved
dividing ridge that runs
through the middle of
the MACULA in the
UTRICLE, the kinocilia
are oriented TOWARD
the striola, and in the
SACCULE they are
oriented AWAY from it
in any position, some
hair cells will be
depolarized and others
hyperpolarized in BOTH
otolith organs

Signal transduction

Semicircular canals- coding of

direction
Three semicircular canals in each ear

Each canal is oriented in a different plane

Each canal is maximally sensitive to rotations

perpendicular to the canal plane

Function of the cristae

detect the rate of head

rotation
when the head is initially
moved, the endolymph
and ampulla (and therefore
the hair cells) turn with it in
a direction oposing rotation
DEPOLARISATION IN
THE KINOCILUM
DIRECTION
HYPERPOLARISATION IN
THE STEREOCILLIUM
DIRECTION

HORIZONTAL CANALS (no)

depolarization

occurs in the
SAME direction as
the head
movement (LEFT
head turn
produces
depolarization in
the LEFT
horizontal canal)

ANTERIOR (SUPERIOR) (YES)

AND POSTERIOR CANALS
anterior canals are located at ~90o to

each other
posterior canals are also located at
~90o to each other
the directionality of the stereocilia is
different in the anterior and posterior
canals
the anterior canals have their

kinocilium anterior to the stereocilia

the posterior canals have their
kinocilium posterior to the
stereocilia
the natural pairing of A/P canals is:

LEFT ANTERIOR with RIGHT

POSTERIOR
RIGHT ANTERIOR with LEFT
POSTERIOR

Signal transduction
K+ channels in the cillia
When stereocilia are bent towards

the kinocilum K+ ch open

Depolarisation of the receptor cell=
receptor potential
Ca2+ channels opening mediator
release in the synaptic cleft
Action potential on the vestibular
pathway

Vestibular pathway
1st order neuron= Scarpa ganglion- axons form the

vestibular branch of the VIIIth cranial nerve

2nd order neuron- vestibular nuclei in the medulla
oblongata
3rd order neuron- inferior colliculus
4th order neuron- medial geniculate nucleus
Cortical projection= superior temporal gyrus

 1. to the

thalamus (3rd
order neuron)
2. to the
paleocerebellum
3. to the spinal
chord
(vestibulospinal)
4. to the motor
nuclei of cranial
nerves III,IV,VI

Chemical senses
Olfaction and taste
Both have receptors that sense the differences of

concentration of a substance dissolved in the mucus/ saliva

So- the odorant/sapide molecules have to be soluble
Interaction of molecules with receptor cells

Both project to cerebral cortex & limbic system

evokes strong emotional reactions

Olfaction

Anatomy of olfactory receptors

The receptors for olfaction, which are bipolar

neurons, are in the nasal epithelium in the superior

portion of the nasal cavity
The only neurons to come into contact with the
external enviroment
They are first-order neurons of the olfactory
pathway.
Basal stem cells produce new olfactory receptors.
The only neurons that regenerate!
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Olfactory Epithelium
1 square inch of membrane

holding 10-100 million receptors

Covers superior nasal cavity and
cribriform plate

24

Cells of the Olfactory Mucosa

Olfactory receptors
bipolar neurons with cilia or

olfactory hairs
Supporting cells
columnar epithelium
Basal cells = stem cells
replace receptors monthly
Olfactory glands (Bowmann)
produce mucus

Principles of Human Anatomy and Physiology, 11e

25

Olfaction: Sense of Smell

Principles of Human Anatomy and Physiology, 11e

Odorants bind to receptors

Na+/ Ca2+/ Cl- channels open
Depolarization occurs

Nerve impulse is triggered

26

Olfactory receptors use a G-protein coupled

transduction mechanism similar to visual receptors

There are 1000 different genes in 4 families; each codes 7transmembrane domain G-protein coupled receptor protein
that is expressed in olfactory receptors in mice

About 350 of these are functional genes in humans; the rest

are present as pseudogenes
Each receptor cell in the epithelium expresses only one
receptor gene
Therefore, each receptor is best tuned to one of 1000
different chemical types
What these types are is still not clear, nor is how the code gets
turned into a smell

RECEPTORS

There are, in the rat, about 1000 odorant receptor genes. Each olfactory receptor expresses
only one of these genes. This is the first critical feature of olfactory coding. When an odorant
binds to the olfactory receptor protein it stimulates a G-protein that activates adenylate
cyclase; cAMP binds to and opens channels permeable to Na+/Ca2+ and Cl- channels. The
resulting current flow depolarizes the receptor cell (receptor potential) causing it to spike. Its
axon terminal in the OB then releases transmitter (glutamate) to excite the target mitral cells.
Bear et al.

OLFACTORY BULB
Bear et al.

Olfactory receptor axons terminate on mitral cell

dendrites in a restricted encapsulated structure
called a glomerulus; a glomerulus contains the
dendritic bush of one mitral cell but many
olfactory receptor axons. All the OR axons ending
in one glomerulus contain are from receptors
So each mitral cell codes for one kind of
expressing same olfactory binding protein.
odorant molecule. This is the primary basis
of olfactory coding.

Olfactory Pathway
Axons from olfactory receptors form the olfactory nerves

(Cranial nerve I) that synapse in the olfactory bulb

pass through 40 foramina in cribriform plate

Second-order neurons within the olfactory bulb form the

olfactory tract that synapses on primary olfactory area of

temporal lobe, in the paleocortex
conscious awareness of smell begins

Other pathways lead to the frontal lobe (Brodmann area

11) where identification of the odor occurs

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Adaptation & Odor Thresholds

Adaptation = decreasing sensitivity
Olfactory adaptation is rapid
50% in 1 second
complete in 1 minute
Low threshold
only a few molecules need to be present
methyl mercaptan added to natural gas as warning

Taste

Taste receptors
Clustered in taste

buds
Associated with
lingual papillae
OR free in the
pharynx
Contain basal cells
which appear to
be stem cells
Gustatory cells
extend taste hairs
through a narrow
taste pore

Gustatory discrimination
Primary taste sensations
Sweet, sour, salty, bitter, umami (aminoacid glutamate)
Taste sensitivity shows significant individual differences,

some of which are inherited

The number of taste buds declines with age

Taste buds
Receptors for taste are

modified epithelial cell

present in taste buds
located on the tongue,
roof of the mouth and
pharynx

Taste receptors

FLAVOR