Physiology of

Special Sense

Ronny
Head of Subdivision Physiology
Dept. Afisel
Faculty of Medicine
Universitas Padjadjaran
The Senses
 General senses of touch (tactile)
 Temperature- thermoreceptors (heat)
 Pressure- mechanoreceptors (movement)
 Pain- mechanoreceptors
 Special senses
 Smell- chemoreceptors (chemicals)
 Taste- chemoreceptors
 Sight- photoreceptors (light)
 Hearing- mechanoreceptors
 Equilibrium- (balance) mechanoreceptors
Audiovestibular System

 Before start this lab activity, student

should know about:
◦ Anatomy of hearing organ
◦ Anatomy of vestibular organ
Introduction
 Sensory Systems
◦ Sense of hearing, audition
 Detect sound
 Perceive and interpret nuances
◦ Sense of balance, vestibular system
 Head and body location
 Head and body movements
AUDITORY SYSTEM
The Nature of Sound

 Sound
◦ Audible variations in air pressure
◦ Sound frequency: Number of cycles per
second expressed in units called Hertz (Hz)
◦ Cycle: Distance between successive
compressed patches
◦ Range: 20 Hz to 20,000 Hz
◦ Pitch: High and Low
◦ Intensity: Difference in pressure between
compressed and rarefied patches
Structure of Auditory System
 Auditory Pathway Stages
 Sound waves
 Tympanic membrane
 Ossicles
 Oval window
 Cochlea fluid
 Sensory neuron response
 The Middle Ear
 Sound Force Amplification by the Ossicles
◦ Pressure: Force by surface area
◦ Greater pressure at oval window than tympanic
membrane, moves fluids
 The Attenuation Reflex
◦ Response where onset of loud sound causes
tensor tympani and stapedius muscle contraction
◦ Function: Adapt ear to loud sounds, understand
speech better
Middle and Inner Ear
Middle ear-auditory ossicles
Figure 16.15
Figure 16.16
Figure 16.17
The Inner Ear
 Anatomy of the Cochlea

 Perilymph: Fluid in scala vestibuli and scala tympani

 Endolymph: Fluid in scala media
 Endocochlear potential: Endolymph electric
potential 80 mV more positive than perilymph
Internal ear
The Inner Ear
 Physiology of the Cochlea
◦ Pressure at oval window, pushes perilymph into
scala vestibuli, round window membrane bulges
out
 The Response of Basilar Membrane to
Sound
◦ Structural properties: Wider at apex, stiffness
decreases from base to apex
 Research: Georg von Békésy
◦ Endolymph movement bends basilar membrane
near base, wave moves towards apex
The Inner Ear
 The Organ of Corti and Associated
Structures
The Inner Ear
 Transduction by
Hair Cells
◦ Research: A.J.
Hudspeth.
◦ Sound: Basilar
membrane
upward, reticular
lamina up and
stereocilia bends
outward
The Inner Ear
 The Innervation of Hair Cells
◦ One spiral ganglion fiber: One inner hair cell,
numerous outer hair cells
 Amplification by Outer Hair Cells
◦ Function: Sound transduction
◦ Motor proteins: Change length of outer hair
cells
◦ Prestin: Required for outer hair cell
movements
Central Auditory Processes
 Auditory Pathway
◦ More synapses at nuclei than visual pathway,
more alternative pathways
◦ Anatomy
 Dorsal cochlear nucleus, ventral cochlear nucleus,
superior olive, inferior colliculus, MGN, lateral
lemniscus, auditory nerve fiber
 Primary pathway:Ventral cochlear nucleus to superior
olive to inferior colliculus to MGN to auditory cortex
 Central Auditory Processes
 Response Properties of Neurons in
Auditory Pathway
◦ Characteristic frequency
 Frequency at which neuron is most responsive
◦ Response
 More complex and diverse on ascending auditory
pathway in brain stem
 Encoding Sound Intensity and
Frequency
 Encoding Information About Sound Intensity
◦ Firing rates of neurons
◦ Number of active neurons
 Stimulus Frequency, Tonotopy, Phase Locking
◦ Frequency sensitivity: Basilar membrane
◦ Frequency: Highest at base, lowest at cochlea apex
◦ Tonotopy: Systematic organization of characteristic
frequency within auditory structure
Encoding Sound Intensity and
Frequency
 Phase Locking
◦ Consistent firing of cell at same sound wave
phase

Neuroscience: Exploring the Brain, 3rd Ed,

Bear, Connors, and Paradiso Copyright ©
Slide 25 2007 Lippincott Williams & Wilkins
 Mechanisms of Sound Localization
 Techniques for Sound Localization
◦ Horizontal: Left-right, Vertical: Up-down
 Localization of Sound in Horizontal Plane
◦ Interaural time delay: Time taken for sound to
reach from ear to ear
◦ Interaural intensity difference: Sound at high
frequency from one side of ear
◦ Duplex theory of sound localization:
 Interaural time delay: 20-2000 Hz
 Interaural intensity difference: 2000-20000 Hz
Mechanisms of Sound Localization
The Sensitivity of Binaural Neurons to
Sound Location
Monaural: Sound in one ear
Binaural: Sound at both ears
Superior olive: Cochlear nuclei input to
superior olive, greatest response to specific
interaural delay
Mechanisms of Sound Localization
 Delay Lines and Neuronal Sensitivity to
Interaural Delay
◦ Sound from left side, activity in left cochlear
nucleus, sent to superior olive
◦ Sound reaches right ear, activity in right
cochlear nucleus, first impulse far
◦ Impulses reach olivary neuron at the same
time summation action potential
 Localization of Sound in Vertical Plane
◦ Sweeping curves of outer ear
Auditory Cortex
 Acoustic Radiation
◦ Axons leaving MGN project to auditory cortex
via internal capsule in an array
◦ Structure of A1 and secondary auditory areas:
Similar to corresponding visual cortex areas
 Neuronal Response Properties
◦ Frequency tuning: Similar characteristic
frequency
◦ Isofrequency bands: Similar characteristic
frequency, diversity among cells
Auditory Cortex
 Principles in Study of Auditory Cortex
◦ Tonotopy, columnar organization of cells with
similar binaural interaction
 The Effects of Auditory Cortical Lesions
and Ablation
◦ Lesion in auditory cortex: Normal auditory
function
◦ Lesion in striate cortex: Complete blindness
in one visual hemifield
◦ Different frequency band information: Parallel
processing, localization deficit
VESTIBULAR SYSTEM
 The Vestibular System
 Importance of Vestibular System
◦ Balance, equilibrium, posture, head, body, eye
movement
 The Vestibular Labyrinth
 Lateral line organs
◦ Small pits or tubes
 Function
◦ Sense vibration or
pressure changes
 The Vestibular System
 The Otolith Organs
The Vestibular System
 The Semicircular Canals
◦ Function: Detect head movements
 Structure
◦ Crista: Sheet of cells where hair cells of
semicircular canals clustered
◦ Ampulla: Bulge along canal, contains crista
◦ Cilia: Project into gelatinous cupula
◦ Kinocili oriented in same direction so all excited
or inhibited together
◦ Semicircular canals: Filled with endolymph
The Vestibular System
 Push-Pull Activation of Semicircular
Canals
◦ Three semicircular canals on one side
 Helps sense all possible head-rotation angles
◦ Canal: Each paired with another on opposite
side of head
◦ Push-pull arrangement of vestibular axons:
Rotation causes excitation on one side,
inhibition on the other
The Vestibular System
 Central Vestibular Pathways
The Vestibular System
 The Vestibulo-Ocular Reflex (VOR)
◦ Function: Line of sight fixed on visual target
◦ Mechanism: Senses rotations of head,
commands compensatory movement of eyes
in opposite direction
◦ Connections from semicircular canals, to
vestibular nucleus, to cranial nerve nuclei 
excite extraocular muscles
The Vestibular System
 Vestibular Pathology
◦ Drugs can damage vestibular system
◦ Effects:
 Trouble fixating on visual targets
 Walking and standing difficult
Figure 16.19
Figure 16.19a
Figure 16.19b
Figure 16.19c
Figure 16.20
Figure 16.20a
Figure 16.20b
Figure 16.20c
Figure 16.21
Maculae-structure and receptor localization
Semicircular ducts
Figure 16.18
Figure 16.18b
Concluding Remarks
 Hearing and Balance
◦ Nearly identical sensory receptors (hair cells)
◦ Movement detectors: Periodic waves,
rotational, and linear force
◦ Auditory system: Senses external
environment
◦ Vestibular system: Senses movements of itself
Concluding Remarks
 Hearing and Balance (Cont’d)
◦ Auditory Parallels Visual System
 Tonotopy (auditory) and Retinotopy (visual)
preserved from sensory cells to cortex code
◦ Convergence of inputs from lower levels 
Neurons at higher levels have more complex
responses
Lab Activity
 Auditory System
◦ Rinne Test
◦ Weber Test
 Vestibular System
◦ Barany Chair
Rinne test
 strike tuning fork and place on mastoid process
◦ physicians typically use tuning forks of either 256 or
512 Hz (i.e., low frequencies of human hearing), which
means that they are testing the stiffness component
of the middle ear's acoustic impedance, which is
determined by the volume of the middle ear space
 when person can no longer hear the
sound (bone conduction threshold), place
the tuning fork near the ear canal
 ask when the person can no longer hear
the sound (air conduction threshold)
 due to the amplification provided by the
middle ear, in a normal person air
threshold <= bone threshold
 this is usually stated as air conduction is
better than bone conduction (AC > BC)
 in a person with a conductive hearing loss, bone
conduction is greater than air conduction (BC >
AC; bone threshold is less than air threshold)
 in a patient with a profound,
unilateral sensorineural hearing loss, there may be
a false negative Rinne due to the response of
the normal ear i.e., AC>BC in the affected ear, but
both of these thresholds are elevated relative to
BC in the normal ear, so the patient reports that
bone conduction is louder (they're actually
hearing the tone in their normal ear, rather than
their tested --- i.e., affected --- ear)
 Weber Test
◦ Performed with 512 or 1024 tuning forks
◦ Compares patient perception of loudness of air
conduction versus bone condition
◦ Turning fork is struck and base placed on either
the forehead, vertex or upper incisor teeth
◦ Patient is asked where the sound is heard the
loudest.
◦ Normal hearing person: Sound is heard equally in
both eyes.
 Weber Test
Vestibular System
 Barany test
◦ THE BARANY CHAIR, NAMED AFTER THE
AUSTRO-HUNGARIAN OTOLOGIST RÓBERT
BÁRÁNY, IS A DEVICE WHICH IS USED FOR
AEROSPACE PHYSIOLOGY TRAINING, ESPECIALLY
FOR STUDENT PILOTS.
◦ FOR HIS WORK ON THE PHYSIOLOGY AND
PATHOLOGY OF THE VESTIBULAR APPARATUS
BÁRÁNY RECEIVED THE 1914 NOBEL PRIZE IN
PHYSIOLOGY OR MEDICINE.
What to do?
 Nystagmus
 Past pointing test of Barany
 Fall down test
 Sensation
Nystagmus test
 Sit on Barany Chair with closed eyes,
make angle.30° to the front.
 Circle 10 times in 20 sec
 Stop the chair
 Open the eyes
 Check the nystagmus
 Report the results and underlying
mechanism of this phenomenon!
Past point test of Barany
 Sit on Barany Chair with closed eyes,
make angle.30° to the front.
 Circle 10 times in 20 sec
 Stop the chair
 Open the eyes
 Ask the person to pointing the
examiner`s pointing finger
 Report the results and underlying
mechanism of this phenomenon
Fall down test
 Sit on the Barany chair with closed
eyes. Sit with angle 120°. WHAT
THE PURPOSE?
 Circle 10x in 10 sec regularly
 Stop the chair
 Ask the person to sit straightly
 Note the fall down direction
 Report the results and underlying
mechanism of this phenomenon!
Sensation test
 Use the other person
 Sit straight on Barany chair with closed eyes
 Circle the barany chair to right direction with
acceleration speed. Put the steady speed for 5 sec
then apply deceleration speed until stop.
 Note what the person feel when:
◦ Acceleration speed
◦ Steady speed
◦ Deceleration speed
◦ Stop
 Report the results and underlying
mechanism of this phenomenon
Review
 Write the results of Audiometry test,
Rinne and Weber
 Write the results and underlying
mechanism of Vestibular test,
◦ Nystagmus test
◦ Past pointing test of Barany
◦ Fall down test
◦ Sensation test
Visual System
 Eye
 Accessory structures
◦ Eyebrows, eyelids, eyelashes, tear glands
◦ Protect eyes from sunlight and damaging particles
 Optic nerve (II)
◦ Tracts
◦ Pathways
 Eyes respond to light and initiate afferent action potentials
Accessory Structures of Eye
 Eyebrows
◦ Prevent running perspiration
into eyes
◦ Shade
 Eyelids or palpebrae
◦ Consist of 5 tissue layers
◦ Protect and lubricate
 Conjunctiva
◦ Covers inner eyelid and
anterior part of eye
 Lacrimal apparatus
 Extrinsic eye muscles
 Lacrimal Apparatus
1. Lacrimal Gland: Produces
tears to moisten, lubricate,
wash. Innervated by
parasympathetic fibers of facial
(VII) nerve.
Production=1ml/day
2. Lacrimal Canaliculi
◦ Collects excess tears
3. Punctum-small openings
4. Lacrimal Sac
5. Nasolacrimal duct
◦ Opens into nasal cavity

Flow of tears: Lacrimal gland--excretory lacrimal ducts--sup. or inf. lacrimal canal--

lacrimal sac--nasolacrimal duct--nasal cavity.
Extrinsic Eye Muscles
Anatomy of the Eyeball
Anterior and Posterior chambers of the Eye
 Eyeball Anatomy

Cornea- tranparent coat curved to help focus light onto retina.

Sclera-the white of the eye made of collagen fibers and fibroblasts

Uvea-
choroid-lines the sclera
ciliary body-appears dark brown because of melanocytes
Iris-colored portion of the eyeball, contains melanocytes: brown-high melanin,
green-medium melanin, blue-low melanin.

Retina-beginning of the visual pathway contains photoreceptor cells.

Responses of the pupil to light
Retina

Exit of
optic nerve
 The Retina
Four kinds of light-sensitive receptors are found in the retina:
･rods
･three kinds of cones, each "tuned" to absorb light from a portion of the spectrum of visible
light
･cones that absorb long-wavelength light (red)
･cones that absorb middle-wavelength light (green)
･cones that absorb short-wavelength light (blue)

Each type of receptor has its own special pigment for absorbing light. Each consists of:･a
transmembrane protein called opsin coupled to･the prosthetic group retinal. Retinal is a
derivative of vitamin A (which explains why night blindness is one sign of vitamin A deficiency)
and is used by all four types of receptors.

The retina also contains a complex array of interneurons:･bipolar cells and ganglion cells
that together form a path from the rods and cones to the brain･a complex array of other
interneurons that form synapses with the bipolar and ganglion cells and modify their
activity.Ganglion cells are always active. Even in the dark they generate trains of action potentials
and conduct them back to the brain along the optic nerve.Vision is based on the modulation of
these nerve impulses. There is not the direct relationship between visual stimulus and an action
potential that is found in the senses of hearing, taste, and smell. In fact, action potentials are not
even generated in the rods and cones.
 Photoreceptors
Two type of photoreceptors:
1. Rods- function only in dim light and are blind to
color.
2. Cones- operate in bright light and are responsible
for high acuity vision, as well as color.

Rods and cones form an uneven mosaic within the

retina, with rods generally outnumbering cones
more than 10 to 1 except in the retina's center, or
fovea.The cones are highly concentrated in the
fovea. Even though the fovea is essential for fine
vision, it is less sensitive to light than the
surrounding retina. Thus, if we wish to detect a
faint star at night, we must gaze slightly to the side
of the star in order to project its image onto the
more sensitive rods, as the star casts insufficient
light to trigger a cone into action.
 Photopigment
Rods-rhodopsin
Cones- 3 types

Photopigments:
1. Opsin-glycoprotein
2. Retinal-light absorbing derivative of
vitamin A (formed from carotene-
”eat carrots for good vision”)
Rod photoreceptors
The lens
The lens is located just behind the iris. It is held in position by zonules extending from an
encircling ring of muscle. When this ciliary muscle is･relaxed, its diameter increases, the
zonules are put under tension, and the lens is flattened;･contracted, its diameter is reduced,
the zonules relax, and the lens becomes more spherical. These changes enable the eye to
adjust its focus between far objects and near objects.

Farsightedness. If the eyeball is too short or the lens too flat or inflexible, the light rays
entering the eye particularly those from nearby objects will not be brought to a focus by the
time they strike the retina. Eyeglasses with convex lenses can correct the problem.
Farsightedness is called hypermetropia.

Nearsightedness. If the eyeball is too long or the lens too spherical, the image of distant
objects is brought to a focus in front of the retina and is out of focus again before the light
strikes the retina. Nearby objects can be seen more easily. Eyeglasses with concave lenses
correct this problem by diverging the light rays before they enter the eye. Nearsightedness is
called myopia.
Cataracts One or both lenses often become cloudy as one ages. When a cataract
seriously interferes with seeing, the cloudy lens is easily removed and a plastic one substituted.
The entire process can be done in a few minutes as an outpatient under local anesthesia.

The iris and lens divide the eye into two main chambers･the front chamber is filled with a
watery liquid, the aqueous humor･the rear chamber is filled with a jellylike material, the
vitreous body.
Refraction
Accommodation

Increased curvature of the lens for

near vision
Visual Pathway
Visual Pathways
Olfactory System
 Nose- olfactory epithelium-receptors, supporting cells, and basal cells
 Olfactory Receptors- bipolar neuron, exposed knob shaped dendrite and
axon that projects through the cribiform plate synapsing in the olfactory bulb.
 Supporting cells- columnar epithelium-mucous membrane in the lining of
the nose.
 Basal cells- stem cells underneath the supporting cells that divide to produce
new olfactory receptor cells.
 Olfactory nerve (I)- olfactory receptor axons bundle and form the
olfactory nerve which terminate in the olfactory bulb. Axons of olfactory
bulb form the olfactory tract that projects to the primary olfactory area (inf.
and med. temporal lobe), frontal lobe, limbic system, and hypothalamus.
 Olfactory Physiology
• Olfactory receptors respond to odors and produce a generator potential that
triggers an action potential in olfactory nerves that terminate in the olfactory bulb.

• Odorant--binds to a G-protein linked receptor--biochemical cascade--opening Na+

channels--depolarizing generator potential--generation of action potential.

•Adaptation- Occurs rapidly in olfactory receptors. There is a component of olfactory

perception that adapts more slowly and is likely in the brain.
Olfactory Epithelium and Receptors
Olfaction
 Sense of smell
◦ Olfactory neurons in this
epithelium
 Bipolar neurons
◦ Olfactory hairs
 Cilia which lies in mucous
 Odors
◦ Odorants bind to
chemoreceptor molecules
◦ Depolarize and initiate action
potentials in neurons
◦ Low threshold for odor
detection
Figure 16.8
Neuronal Pathways of Olfaction
 Taste
 Sense is detected by taste
buds
 Papillae
◦ Vallate- 12 form an inverted V at  Function
back of tongue-contain 100-300 ◦ Receptors on hairs detect
taste buds.
◦ Fungiform- mushroom shaped all dissolved substances
over tongue each has 5 taste buds. ◦ Adaptation to taste is rapid
◦ Foliate- lateral tongue, but variable based on taste
degenerate in childhood type. Taste aversion-type of
◦ Filiform- pointed structures with adaptation
no taste buds, help tongue to move
food in mouth (friction).  Taste types
 Histology
◦ Sour
◦ Support cells
◦ Gustatory cells ◦ Salty
 Hairs ◦ Bitter
◦ Sweet
◦ Umami (meaty, MSG)
 Taste Receptors
The sense of taste is mediated by taste receptor cells which are bundled in clusters called
taste buds. Taste receptor cells sample oral concentrations of a large number of small molecules
and report a sensation of taste to centers in the brainstem. In most animals, including humans,
taste buds are most prevalent on small pegs of epithelium on the tongue called papillae.

When taste cells are stimulated by binding of chemicals to their receptors, they depolarize and
this depolarization is transmitted to the taste nerve fibers resulting in an action potential that is
ultimately transmitted to the brain. One interesting aspect of this nerve transmission is that it rapidly
adapts - after the initial stimulus, a strong discharge is seen in the taste nerve fibers but within a
few seconds, that response diminishes to a steady-state level of much lower amplitude.
Gustatory (taste) Receptors
Papillae and Taste Buds
Actions of Major Tastants
 Neuronal Pathways for Taste

Parietal lobe

Front 2/3 of tongue

Rear 1/3 of tongue

Throat and epipglottis
Physiology of Gustation

1. Tastant dissolved in saliva

2. binds to gustatory hairs generating a receptor potential that
3. stimulates neurotransmitter secretion
4. Neurotransmitters bind to receptors on the first order sensory neurons that are
postsynaptic to the gustatory receptor cells.

• Receptor potential arises differently for different tastes.

1. Salty taste-Na+ enters the gustatory receptor cells through channels,
causes depolarization, opens Ca2+ channels and triggers neurotransmitter secretion.
2. Sour taste-H+ ions entering H+ channels influencing other channels
resulting in depolarization and neurotransmitter secretion.
3. Sweet, bitter, and umami tastes- tastants bind to G-protein linked
receptors that result in depolarization of the receptor cells and ultimately
neurotransmitter secretion.