Week 3b

Special Senses (Chapter 15)

• Special senses of body include:
– Vision
– Taste
– Smell
– Hearing
– Equilibrium

• All use special sensory receptors, which are distinct receptor

cells localized in head region
– Not like the modified nerves of general receptors
1
 The Eye and Vision
• 70% of body’s sensory receptors are in eye
• Half of cerebral cortex is involved in visual processing

The Eye
• Small sphere; only one-sixth of surface visible
• Most of eye enclosed and protected by fat cushion and bony
orbit
• Consists of accessory structures and the eyeball

2
 Structure of the Eyeball
• Wall of eyeball
contains three layers
– Fibrous layer
(sclera and
cornea)
– Vascular layer
(choroid, ciliary
body, iris)
– Inner layer (retina)
• Lens separates internal
cavity into anterior and
posterior segments
• Internal cavity filled
with fluids called
humors Fig 15.4 3
Fig 15.6 4
Neural layer of the retina
§ Transparent layer
§ Composed of three main
types of neurons
• Photoreceptors, bipolar
cells, ganglion cells
§ Signals spread from
photoreceptors to bipolar
cells to ganglion cells
§ Ganglion cell axons exit eye
as optic nerve
§ Retina has quarter-billion
photoreceptors that are one
of two types:
– Rods
– Cones Fig 15.6
5
Photoreceptors
- Rods
§ Dim light, peripheral vision receptors
§ More numerous and more sensitive to light than cones
§ No color vision or sharp images
§ Numbers greatest at periphery

– Cones
§ Vision receptors for bright light
§ High-resolution color vision
§ Macula lutea area at posterior pole lateral to blind spot
– Contains mostly cones
§ Fovea centralis: tiny pit in center of macula lutea that
contains all cones, so is region with best visual acuity
– Eye movement allows us to focus in on object so that
fovea can pick it up 6
Functional Anatomy of Photoreceptors
• Photoreceptors (rods and cones)
are modified neurons
• Consists of cell body, synaptic
terminal, and two segments:
– Outer segment: light-receiving
region
§ Contains visual pigments
(photopigments) that
change shape as they
absorb light
– Inner segment of each joins Fig 15.15

cell body
§ Inner segment is
connected via cilium to
outer segment and to cell 7
body via outer fiber
Functional Anatomy of Photoreceptors
– Plasma membrane of outer Fig 15.15
segment folds to form many
discs
§ Photopigments are
embedded in discs

8
Comparing Rod and Cone Vision
• Rods are very sensitive to light, making them best suited for
night vision and peripheral vision
– Contain a single pigment, so vision is perceived in gray tones
only
– Pathways converge, causing fuzzy, indistinct images
§ As many as 100 rods may converge into one ganglion cell
• Cones have low sensitivity, so require bright light for activation
– React more quickly than rods
– Have one of three pigments (blue, red or green), which
allow for vividly colored sight
– Nonconverging pathways result in detailed, high-resolution
vision
§ Some cones have their own ganglion cell, so brain can
put together accurate, high-acuity resolution images
9
Visual Pigments
• Retinal: key light-absorbing molecule that combines with one of
four proteins (opsins) to form visual pigments
– Four opsins are rhodopsin (found in rods only), and three
found in cones: green, blue, red (depending on wavelength
of light they absorb)
– Retinal isomers are different 3-D forms
§ Retinal is in a bent form in dark, but when pigment
absorbs light, it straightens out
– Bent form called 11-cis-retinal
– Straight form called all-trans-retinal
§ Conversion of bent to straight initiates reactions that
lead to electrical impulses along optic nerve

10
Fig 15.16

11
Phototransduction
• Phototransduction: process by which pigment captures photon
of light energy, which is converted into a graded receptor
potential

• Light transduction reactions

– Light-activated rhodopsin activates G protein transducin
– Transducin activates Phosphodiesterase, which breaks down
cyclic GMP (cGMP)
– In dark, cGMP holds ion channels of outer segment open
§ Na+ and Ca2+ enter and depolarize cell to ~-40mV
– With light exposure the activated phosphodiesterase breaks
down cGMP, ion channels close, cell hyperpolarizes to -
70mV
§ Hyperpolarization is signal for vision!
12
 13
Figure 15.17 Events of phototransduction.
Information Processing in the Retina

• Photoreceptors and bipolar cells generate only graded

potentials (EPSPs and IPSPs), not APs
• When light hyperpolarizes photoreceptor cells, they stop
releasing inhibitory neurotransmitter glutamate to bipolar cells
• Bipolar cells (no longer inhibited) depolarize, release
neurotransmitter onto ganglion cells
• Ganglion cells generate APs transmitted in optic nerve to brain

14
Fig 15.18

15
Visual Pathway to the Brain
• Axons of retinal ganglion cells form optic nerve
• Medial fibers from each eye cross over at the optic chiasma
then continue on as optic tracts, which means each optic tract:
– Contains fibers from lateral (temporal) aspect of eye on
same side and medial (nasal) aspect of opposite eye, and
– Each carries information from same half of visual field
• Most fibers of optic tracts continue on to the thalamus
• From there, thalamic neurons form optic radiation, which
projects to primary visual cortex in occipital lobes
– Conscious perception of visual images occurs here

16
 Visual Pathway to the Brain and Visual Fields, Inferior View
• A small subset of ganglion cells
in retina contains melanopsin
(circadian pigment), which
project to:
– Suprachiasmatic nucleus
of hypothalamus: timer for
daily biorhythms
– Pretectal nucleus –
mediate pupillary light
reflexes

Figure 15.19a
17
The Chemical Senses: Smell and Taste
• Smell (olfaction) and taste (gustation): complementary senses
that let us know whether a substance should be savored or
avoided
• Chemoreceptors are used by these systems
– Chemicals must be dissolved in aqueous solution to be
picked up by chemoreceptors
§ Smell receptors are excited by chemicals dissolved in
nasal fluids
§ Taste receptors respond to chemicals dissolved in saliva

18
 Location and Structure of Olfactory Receptors
• Olfactory epithelium: organ of
smell
– Located in in roof of nasal
cavity
– Covers superior nasal
conchae

Fig 15.20

19
• Contains olfactory
sensory neurons
- Bipolar neurons
with radiating
olfactory cilia
• Supporting cells
surround and
cushion olfactory
receptor cells
• Olfactory stem cells
lie at base of
epithelium
• Axons of sensory
neurons are grouped
together and called
filaments of olfactory
nerve 20
Fig 15.20
Specificity of Olfactory Receptors
• Smells may contain 100s of different chemicals (odorants)
• Humans have ~400 “smell” genes active in nose
– Each encodes a unique receptor protein
§ Protein responds to one or more odorants
– Each odorant binds to several different receptor proteins
– Each bipolar neuron has one type of odorant receptor
protein
• Pain and temperature receptors are also in nasal cavities
– Respond to irritants, such as ammonia, or can “smell” hot
or cold (chili peppers, menthol)

21
Physiology of Smell
• In order to smell substance, it must be volatile
– Must be in gaseous state
– Odorant must also be able to dissolve in olfactory
epithelium fluid
• Activation of olfactory sensory neurons
– Dissolved odorants bind to receptor proteins in olfactory
cilium membranes
§ Open ion channels, generating receptor potential
§ At threshold, AP is conducted to first relay station in
olfactory bulb

22
• Smell transduction
– Odorant binds to receptor, activating a Golf protein
– G protein activation causes cAMP (second messenger) synthesis
– cAMP opens Na+ and Ca2+ channels
– Na+ influx causes depolarization and impulse transmission
– Ca2+ influx causes decreased response to a sustained
stimulus, referred to as olfactory adaptation
§ People can’t smell a certain odor after being exposed to
it for a while
Fig 15.21

23
The Olfactory Pathway
• Filaments of olfactory nerves (axons of olfactory sensory neurons)
synapse with mitral cells located in overlying olfactory bulb
– Mitral cells are second-order neurons that form olfactory tract
• Synapse occurs in structures called glomeruli (see Fig 15.20)
• Mitral cells amplify, refine, and relay signals
• Impulses from activated mitral cells travel via olfactory tracts (axons
of mitral cells) to lobe of primary olfactory cortex
• Some information sent to frontal lobe, and some passes through
thalamus first
– Smell is consciously interpreted and identified
• Some information sent to hypothalamus, amygdala, and other
regions of limbic system
– Emotional responses to odor are elicited
24
 The Ear: Hearing and Balance
• The ear has three major
areas:
Fig 15.24
– External (outer) ear:
hearing only
– Middle ear (tympanic
cavity): hearing only,
3 auditory ossicles
– Internal (inner) ear:
receptors for hearing
and equilibrium
• Receptor neurons for
hearing and balance
respond to separate
stimuli and are activated
independently 25
• Auditory ossicles: three small bones in tympanic cavity, named
for their shape:
– Malleus: the “hammer” is secured to eardrum
– Incus: the “anvil”
– Stapes: the “stirrup” base fits into oval window

Fig 15.24 26
Internal Ear (labyrinth)
• Two major divisions:
– Bony labyrinth: system of tortuous channels and cavities that
worm through the bone
– Divided into three regions: vestibule, semicircular canals
(balance), and cochlea (hearing)
§ Filled with perilymph fluid (similar to CSF)
– Membranous labyrinth; series of membranous sacs and ducts
contained in bony labyrinth; filled with potassium-rich endolymph

27
Membranous Labyrinth (blue) of the Internal Ear contained in bony labyrinth
Fig 15.26

28
• Semicircular canals (balance)
– Three canals oriented in three planes of space: anterior, lateral,
and posterior
§ Anterior and posterior are at right angles to each other,
whereas the lateral canal is horizontal
– Ampulla: enlarged area of ducts of each canal that houses
equilibrium receptor region called the crista ampullaris
§ Receptors respond to angular (rotational) movements of the
head
• Vestibule (balance)
– Central egg-shaped cavity of bony labyrinth
– Contains two membranous sacs

29
• Cochlea (hearing)
– A small spiral, conical, bony chamber, size of a split pea
§ Coils around bony pillar Contains cochlear duct, which
houses spiral organ (organ of Corti) and ends at cochlear
apex
• Cavity of cochlea divided into three chambers including cochlear
duct (middle chamber), scala tympani and scala vestibuli
• “Floor” of cochlear duct composed of:
– Bony spiral lamina
– Basilar membrane, which supports spiral organ
• Spiral organ contains cochlear hair cells functionally arranged in
one row of inner hair cells and three rows of outer hair cells
– Hair cells are sandwiched between tectorial and basilar
membranes
• The cochlear branch of nerve runs from spiral organ to brain 30
Figure 15.27a Anatomy of the cochlea.
31
32
 Sound Detection
• Hearing is the reception of an air sound wave that is converted to a
fluid wave in the cochlea that ultimately stimulates mechanosensitive
cochlear hair cells that send impulses to the brain for interpretation
• Pathway of sound (see Fig 15.30)
– Tympanic membrane: sound waves strike tympanic membrane,
causing it to vibrate
– Auditory ossicles: transfer vibration of eardrum to oval window
– Scala vestibuli: vibration of oval window causes wave motions
in liquid perilymph contained in the bony labyrinth
§ Wave ends at round window,
– Helicotrema path: waves with frequencies below threshold of
hearing travel through helicotrema and scali tympani to round
window and don’t excite hair cells
– Basilar membrane path: sounds in hearing range go through
endolymph containing cochlear duct, vibrating basilar
membrane at specific location, according to frequency of sound33
Figure 15.30 Pathway of sound waves.
34
 Resonance of the Basilar Membrane
• Resonance: movement of different areas of
basilar membrane in response to a
particular frequency
• Basilar membrane changes along its length:
– Fibers near oval window are short and
stiff
§ Resonate with high-frequency waves
– Fibers near cochlear apex are longer,
floppier
§ Resonate with lower-frequency
waves
• Impulses from specific hair cells interpreted
as pitch
Fig 15.31
35
Sound Transduction
• Excitation of inner hair cells
– Vibrations within the hearing range of the animal causes
vibration of basilar membrane in the floor of the cochlear
duct which deflects hairs (cilia) of inner hair cells
§ Cochlear hair cells have microvilli that contain many
stereocilia (hairs) that bend at their base
§ Longest hair cells are connected to shortest hair cells via
tip links
– Tip links, when pulled on, open ion channels they are
connected to
– Stereocilia project into K+-rich endolymph, with longest hairs
enmeshed in gel-like tectorial membrane

36
• Excitation of inner hair cells Fig 15.32
– Bending of stereocilia toward tallest ones pull on tip links, causing
K+ and Ca2+ ion channels in shorter stereocilia to open
§ K+ and Ca2+ flow into cell, causing graded receptor potential
that can lead to release of neurotransmitter (glutamate)
– Can trigger AP in afferent neurons of cochlear nerve
– Bending of stereocilia toward shorter ones causes tip links to relax
§ Ion channels close, leading to repolarization

37
 Auditory Pathway
• Neural impulses from cochlear
bipolar cells reach auditory
cortex via following pathway:
– Spiral ganglion ®
– medulla oblongata ®
– midbrain auditory reflex
center ®
– thalamus ®
– Primary auditory cortex

38
 Auditory Processing
• Perception of pitch: impulses from hair cells in different
positions along basilar membrane are interpreted by brain as
specific pitches
• Detection of loudness is determined by brain as an increase in
the number of action potentials (frequency) that result when
hair cells experience larger deflections as a result of sound
waves of higher amplitude

• Localization of sound depends on relative intensity and relative

timing of sound waves reaching both ears
– If timing is increased on one side, brain interprets sound as
coming from that side

39
Maintenance of Equilibrium
• Equilibrium is response to various movements of head that rely
on input from inner ear, eyes, and stretch receptors
• Vestibular apparatus: equilibrium receptors in semicircular
canals and vestibule
– Vestibular
receptors
monitor static
equilibrium
– Semicircular
canal receptors
monitor dynamic
equilibrium
40
• Maculae: sensory receptor Forward/
organs that monitor static backward
equilibrium in saccule and Up/down
utricle walls in vestible
- Monitor the position of
head in space
- Play a key role in control of
posture
- Respond to linear
acceleration forces, but not
rotation
• Hair cells have stereocilia,
including kinocilium
- Stereocilia are embedded in
otolith membrane, jelly-like
mass studded with otoliths
(tiny CaCO3 stones) 41
Fig 15.34
• Anatomy of a macula
(cont.)
– Hair cells synapse with
dendrites (fibers) of
vestibular nerve whose
cell bodies are located
in superior and inferior
vestibular ganglia
§ Part of
vestibulocochlear
cranial nerve

42
The Cristae Ampullares of Semi-Circular Canals
• Receptor for rotational acceleration is crista ampullaris in ampulla
• Cristae are excited by acceleration and deceleration of head
– Major stimuli are rotational (angular) movements, such as
twirling of the body
– Semicircular canals are located in all three planes of space, so
cristae can pick up on all rotational movements of head

43
Fig 15.35
• Anatomy of a crista ampullaris
– Each crista has supporting cells and hair cells that extend
into gel-like mass called ampullary cupula
– Dendrites of vestibular nerve fibers encircle base of hair
cells
• Activating receptors of crista ampullaris
– Cristae respond to changes in velocity of rotational
movements of head
– Inertia in ampullary cupula causes endolymph in
semicircular ducts to move in direction opposite body’s
rotation, causing hair cells to bend
– Axes of hair cells in complementary semicircular ducts are
opposite
§ Depolarization occurs in one ear, while hyperpolarization
occurs in other ear
– Endolymph will come to rest after a while, so this system
detects only changes in movements 44
 45
Fig 15.35
 Equilibrium Pathway to the Brain
• Equilibrium information goes to
reflex centers in brain stem
– Allows fast, reflexive
responses to imbalance so
we don’t fall down
• Impulses from activated
vestibular receptors travel to
either vestibular nuclei in brain
stem or to cerebellum
• Three modes of input for
balance and orientation:
– Vestibular receptors
– Visual receptors
– Somatic receptors
(proprioceptors) 46
 Chapter 15 Textbook Questions

Check your understanding: 2, 3, 4, 8, 9,12, 14, 15, 16, 17, 18, 19, 20, 21,

End of Chapter Review:2, 6, 8, 10, 11, 12, 13, 14, 15, 19, 20, 25, 26,

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