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Chapter 17
The Special Senses
Lecture Presentation by
Lee Ann Frederick
University of Texas at Arlington
© 2015 Pearson Education, Inc.
An Introduction to the Special Senses
•  Five Special Senses
1. 
2. 
3. 
4. 
5. 
Olfaction
Gustation
Vision
Equilibrium
Hearing
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17-1 Smell (Olfaction)
•  Olfactory Organs
•  Provide sense of smell
•  Located in nasal cavity on either side of nasal
septum
•  Made up of two layers
1.  Olfactory epithelium
2.  Lamina propria
© 2015 Pearson Education, Inc.
Figure 17-1a The Olfactory Organs.
Olfactory Pathway to the Cerebrum
Olfactory
epithelium
Olfactory
nerve fibers
(N I)
Olfactory
bulb
Olfactory
tract
Central
nervous
system
Cribriform
plate
Superior
nasal concha
a The olfactory organ on the right
© 2015 Pearson Education, Inc.
side of the nasal septum.
Figure 17-1b The Olfactory Organs.
Basal cell: divides
to replace
worn-out
olfactory
Olfactory
gland
receptor cells
To
olfactory
bulb
Cribriform
plate
Olfactory
nerve fibers
Lamina
propria
Developing
olfactory
receptor cell
Olfactory
receptor cell
Olfactory
epithelium
Supporting cell
Mucous layer
Knob
Substance being smelled
b An olfactory receptor is a modified neuron
© 2015 Pearson Education, Inc.
with multiple cilia-shaped dendrites.
Olfactory dendrites:
surfaces contain
receptor proteins
(see Spotlight
Figure 17–2)
17-1 Smell (Olfaction)
•  Olfactory Glands
•  Secretions coat surfaces of olfactory organs
•  Olfactory Receptors
•  Highly modified neurons
•  Olfactory reception
•  Involves detecting dissolved chemicals as they
interact with odorant-binding proteins
© 2015 Pearson Education, Inc.
17-1 Smell (Olfaction)
•  Olfactory Discrimination
•  Can distinguish thousands of chemical stimuli
•  CNS interprets smells by the pattern of receptor
activity
•  Olfactory Receptor Population
•  Considerable turnover
•  Number of olfactory receptors declines with age
© 2015 Pearson Education, Inc.
Figure 17-2 Olfaction and Gustation (Part 4 of 10).
Stimulus
Olfaction and gustation are special
senses that give us vital information
about our environment. Although the
sensory information is diverse and
complex, each special sense originates
at receptor cells that may be neurons
or specialized receptor cells that
communicate with sensory neurons.
mV
Action
potentials
Dendrites
Specialized
olfactory
neuron
Stimulus
removed
−70 mV
Stimulus
Threshold
Generator potential
0
Time (msec)
© 2015 Pearson Education, Inc.
to CNS
Figure 17-2 Olfaction and Gustation (Part 7 of 10).
Olfactory reception occurs on the surface membranes of
the olfactory dendrites. Odorants—dissolved chemicals
that stimulate olfactory receptors—interact with receptors
called odorant-binding proteins on the membrane surface.
1
The binding of an odorant to its
receptor protein leads to the activation
of adenylyl cyclase, the enzyme that
converts ATP to cyclic AMP (cAMP).
Odorant
molecule
MUCOUS
LAYER
2
In general, odorants are small organic molecules. The
strongest smells are associated with molecules of either
high water or high lipid solubilities. As few as four
odorant molecules can activate an olfactory receptor.
The cAMP opens sodium ion
channels in the plasma
membrane, which then begins
to depolarize.
Closed
sodium
channel
If sufficient depolarization
occurs, an action potential is
triggered in the axon, and the
information is relayed to the CNS.
+
+
+
Depolarized
membrane
Receptor
protein
Inactive
G protein
RECEPTOR
CELL
3
cAMP
Active
G protein
adenylate
ATP cyclase cAMP
© 2015 Pearson Education, Inc.
cAMP
+
+
cAMP
Sodium
ions enter
+
17-2 Taste (Gustation)
•  Gustation
•  Provides information about the foods and liquids
consumed
•  Taste Receptors (Gustatory Receptors)
•  Are distributed on tongue and portions of pharynx
and larynx
•  Clustered into taste buds
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17-2 Taste (Gustation)
•  Taste Buds
•  Associated with epithelial projections (lingual
papillae) on superior surface of tongue
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17-2 Taste (Gustation)
•  Four Types of Lingual Papillae
1.  Filiform papillae
•  Provide friction
•  Do not contain taste buds
2.  Fungiform papillae
•  Contain about five taste buds each
3.  Vallate papillae
•  Contain 100 taste buds each
4.  Foliate papillae
© 2015 Pearson Education, Inc.
Figure 17-3a Gustatory Receptors.
Water receptors
(pharynx)
Umami
a Location of
tongue papillae.
© 2015 Pearson Education, Inc.
17-2 Taste (Gustation)
•  Gustatory Discrimination
•  Four primary taste sensations
1. 
2. 
3. 
4. 
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Sweet
Salty
Sour
Bitter
17-2 Taste (Gustation)
•  Additional Human Taste Sensations
•  Umami
•  Characteristic of beef/chicken broths and
Parmesan cheese
•  Receptors sensitive to amino acids, small peptides,
and nucleotides
•  Water
•  Detected by water receptors in the pharynx
© 2015 Pearson Education, Inc.
17-2 Taste (Gustation)
•  Gustatory Discrimination
•  Dissolved chemicals contact taste hairs
•  Bind to receptor proteins of gustatory cell
•  Salt and sour receptors
•  Chemically gated ion channels
•  Stimulation produces depolarization of cell
•  Sweet, bitter, and umami stimuli
•  G proteins
•  Gustducins
© 2015 Pearson Education, Inc.
17-2 Taste (Gustation)
•  End Result of Taste Receptor Stimulation
•  Release of neurotransmitters by receptor cell
•  Dendrites of sensory afferents wrapped by receptor
membrane
•  Neurotransmitters generate action potentials in
afferent fiber
© 2015 Pearson Education, Inc.
17-2 Taste (Gustation)
•  Taste Sensitivity
•  Exhibits significant individual differences
•  Some conditions are inherited
•  For example, phenylthiocarbamide (PTC)
•  70 percent of Caucasians taste it but 30 percent do
not
•  Number of taste buds
•  Begins declining rapidly at age 50
© 2015 Pearson Education, Inc.
Figure 17-2 Olfaction and Gustation (Part 6 of 10).
RECEPTOR CELL
Stimulus
+30
mV
0
Stimulus
removed
Stimulus
Threshold
−60
−70
Receptor
cell
−70 mV
Depolarization
Synapse at
dendrite
0
Time (msec)
Sensory
neuron
−70 mV
AXON
Stimulus
mV
+30
Action
potentials
Synaptic
delay
0
−60
−70
Generator potential
0
© 2015 Pearson Education, Inc.
Time (msec)
to CNS
Figure 17-2 Olfaction and Gustation (Part 10 of 10).
Salt and Sour Channels
Sweet, Bitter, and Umami Receptors
The diffusion of sodium ions from
salt solutions or hydrogen ions from
acids or sour solutions into the
receptor cell leads to depolarization.
Receptors responding to stimuli that
produce sweet, bitter, and umami sensations
are linked to G proteins called gustducins
(GUST-doos-inz)—protein complexes that
use second messengers to produce their effects.
+
H+
+
Na+ +
+
+
Na+ ion
leak channel
Sweet,
bitter, or
umami
Membrane
receptor
Resting plasma
membrane
Inactive
G protein
Depolarized
membrane
+
Depolarization of membrane stimulates
release of chemical neurotransmitters.
© 2015 Pearson Education, Inc.
Depolarized
membrane
Active
G protein
+
+
Active
G protein
Active
2nd messenger
Inactive
2nd messenger
Activation of second messengers stimulates
release of chemical neurotransmitters.
17-3 Accessory Structures of the Eye
•  Accessory Structures of the Eye
•  Provide protection, lubrication, and support
•  Include:
•  The palpebrae (eyelids)
•  The superficial epithelium of eye
•  The lacrimal apparatus
© 2015 Pearson Education, Inc.
Figure 17-4a External Features and Accessory Structures of the Eye.
Eyelashes
Pupil
Lateral
canthus
Palpebra
Palpebral fissure
Sclera
Medial canthus
Lacrimal caruncle
Corneal limbus
a Gross and superficial anatomy
of the accessory structures
© 2015 Pearson Education, Inc.
17-3 Accessory Structures of the Eye
•  Lacrimal Apparatus
•  Produces, distributes, and removes tears
•  Fornix
•  Pocket where palpebral conjunctiva joins ocular
conjunctiva
•  Lacrimal gland (tear gland)
•  Secretions contain lysozyme, an antibacterial
enzyme
© 2015 Pearson Education, Inc.
Figure 17-4b External Features and Accessory Structures of the Eye.
Superior
rectus muscle
Tendon of superior
oblique muscle
Lacrimal
gland ducts
Lacrimal punctum
Lacrimal gland
Lacrimal caruncle
Ocular conjunctiva
Superior lacrimal
canaliculus
Lateral canthus
Medial canthus
Lower eyelid
Inferior lacrimal
canaliculus
Orbital fat
Lacrimal sac
Inferior
rectus muscle
Nasolacrimal duct
Inferior nasal
concha
Inferior
oblique muscle
b The organization of the
lacrimal apparatus
© 2015 Pearson Education, Inc.
Opening of
nasolacrimal duct
Figure 17-5a The Sectional Anatomy of the Eye.
Fornix
Palpebral conjunctiva
Eyelash
Optic
nerve
Ocular conjunctiva
Ora serrata
Cornea
Lens
Pupil
Iris
Corneal
limbus
Fovea
Retina
Choroid
Sclera
a
Sagittal section of left eye
© 2015 Pearson Education, Inc.
Figure 17-5b The Sectional Anatomy of the Eye.
Fibrous
layer
Cornea
Anterior
cavity
Sclera
Vascular layer
(uvea)
Iris
Ciliary body
Choroid
Posterior
cavity
Inner layer
(retina)
Neural part
Pigmented part
b
© 2015 Pearson Education, Inc.
Horizontal section of right eye
Figure 17-5c The Sectional Anatomy of the Eye.
Visual axis
Anterior cavity
Posterior
chamber
Anterior
chamber
Cornea
Edge of
pupil
Iris
Ciliary zonule
Nose
Corneal limbus
Lacrimal punctum
Conjunctiva
Lacrimal caruncle
Lower eyelid
Medial canthus
Ciliary
processes
Lateral
canthus
Lens
Ciliary body
Ora serrata
Sclera
Choroid
Retina
Posterior
cavity
Ethmoidal
labyrinth
Lateral rectus
muscle
Medial rectus
muscle
Optic disc
Fovea
Optic nerve
Central artery
and vein
© 2015 Pearson Education, Inc.
Orbital fat
c
Horizontal dissection of right eye
17-3 The Eye
•  Vascular Layer (Uvea) Functions
1.  Provides route for blood vessels and lymphatics
that supply tissues of eye
2.  Regulates amount of light entering eye
3.  Secretes and reabsorbs aqueous humor that
circulates within chambers of eye
4.  Controls shape of lens, which is essential to
focusing
© 2015 Pearson Education, Inc.
Figure 17-5c The Sectional Anatomy of the Eye.
Visual axis
Anterior cavity
Posterior
chamber
Anterior
chamber
Cornea
Edge of
pupil
Iris
Ciliary zonule
Nose
Corneal limbus
Lacrimal punctum
Conjunctiva
Lacrimal caruncle
Lower eyelid
Medial canthus
Ciliary
processes
Lateral
canthus
Lens
Ciliary body
Ora serrata
Sclera
Choroid
Retina
Posterior
cavity
Ethmoidal
labyrinth
Lateral rectus
muscle
Medial rectus
muscle
Optic disc
Fovea
Optic nerve
Central artery
and vein
© 2015 Pearson Education, Inc.
Orbital fat
c
Horizontal dissection of right eye
17-3 The Eye
•  The Vascular Layer
•  Iris
•  Contains papillary muscles
•  Change diameter of pupil
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Figure 17-6 The Pupillary Muscles.
Pupillary constrictor
(sphincter)
Pupil
The pupillary dilator
muscles extend radially
away from the edge of the pupil.
Contraction of these muscles
enlarges the pupil.
Pupillary dilator
(radial)
Decreased light intensity
Increased sympathetic stimulation
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The pupillary constrictor
muscles form a series of
concentric circles around the
pupil. When these sphincter
muscles contract, the diameter
of the pupil decreases.
Increased light intensity
Increased parasympathetic stimulation
17-3 The Eye
•  The Inner Layer
•  Outer layer called pigmented part
•  Inner called neural part (retina)
•  Contains visual receptors and associated neurons
•  Rods and cones are types of photoreceptors
•  Rods
•  Do not discriminate light colors
•  Highly sensitive to light
•  Cones
•  Provide color vision
•  Densely clustered in fovea, at center of macula
© 2015 Pearson Education, Inc.
Figure 17-5c The Sectional Anatomy of the Eye.
Visual axis
Anterior cavity
Posterior
chamber
Anterior
chamber
Cornea
Edge of
pupil
Iris
Ciliary zonule
Nose
Corneal limbus
Lacrimal punctum
Conjunctiva
Lacrimal caruncle
Lower eyelid
Medial canthus
Ciliary
processes
Lateral
canthus
Lens
Ciliary body
Ora serrata
Sclera
Choroid
Retina
Posterior
cavity
Ethmoidal
labyrinth
Lateral rectus
muscle
Medial rectus
muscle
Optic disc
Fovea
Optic nerve
Central artery
and vein
© 2015 Pearson Education, Inc.
Orbital fat
c
Horizontal dissection of right eye
Figure 17-7a The Organization of the Retina (Part 1 of 2).
Horizontal cell
Cone
Rod
Pigmented
part of retina
Rods and
cones
Amacrine
cell
Bipolar cells
Ganglion cells
LIGHT
a
© 2015 Pearson Education, Inc.
The cellular organization of the retina. The photoreceptors
are closest to the choroid, rather than near the posterior
cavity (vitreous chamber).
Figure 17-7b The Organization of the Retina.
Pigmented
part of retina
Neural part
of retina
Central retinal vein
Optic disc
Central retinal artery
Sclera
Optic nerve
Choroid
b The optic disc in diagrammatic sagittal section.
© 2015 Pearson Education, Inc.
Figure 17-7c The Organization of the Retina.
Fovea
Macula
c
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Optic disc
(blind spot)
Central retinal artery and vein
emerging from center of optic disc
A photograph of the retina as seen through the pupil.
Figure 17-8 A Demonstration of the Presence of a Blind Spot.
© 2015 Pearson Education, Inc.
17-3 The Eye
•  Aqueous Humor
•  Fluid circulates within eye
•  Diffuses through walls of anterior chamber into
scleral venous sinus (canal of Schlemm)
•  Reenters circulation
•  Intraocular Pressure
•  Fluid pressure in aqueous humor
•  Helps retain eye shape
© 2015 Pearson Education, Inc.
Figure 17-9 The Circulation of Aqueous Humor.
Cornea
Anterior cavity
Pupil
Anterior chamber
Scleral venous sinus
Posterior chamber
Body of iris
Ciliary process
Lens
Ciliary zonule
Pigmented
epithelium
Conjunctiva
Ciliary body
Sclera
Posterior cavity
(vitreous chamber)
Choroid
Retina
© 2015 Pearson Education, Inc.
17-3 The Eye
•  The Lens
•  Lens fibers
•  Cells in interior of lens
•  No nuclei or organelles
•  Filled with crystallins, which provide clarity and
focusing power to lens
•  Cataract
•  Condition in which lens has lost its transparency
© 2015 Pearson Education, Inc.
17-3 The Eye
•  Light Refraction
•  Bending of light by cornea and lens
•  Focal point
•  Specific point of intersection on retina
•  Focal distance
•  Distance between center of lens and focal point
© 2015 Pearson Education, Inc.
Figure 17-10 Factors Affecting Focal Distance.
Focal distance
Focal distance
Close
source
Light
from
distant
source
(object)
Focal distance
Focal
point
Lens
a
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The closer the light source,
the longer the focal distance
b
The rounder the lens,
the shorter the focal distance
c
17-3 The Eye
•  Light Refraction of Lens
•  Accommodation
•  Shape of lens changes to focus image on retina
•  Astigmatism
•  Condition where light passing through cornea and
lens is not refracted properly
•  Visual image is distorted
© 2015 Pearson Education, Inc.
Figure 17-11 Accommodation.
a For Close Vision: Ciliary Muscle Contracted, Lens Rounded
Lens rounded
Focal point
on fovea
Ciliary muscle
contracted
b For Distant Vision: Ciliary Muscle Relaxed, Lens Flattened
Lens flattened
Ciliary muscle
relaxed
© 2015 Pearson Education, Inc.
Figure 17-11a Accommodation.
a
For Close Vision: Ciliary Muscle Contracted, Lens Rounded
Lens rounded
Focal point
on fovea
Ciliary muscle
contracted
© 2015 Pearson Education, Inc.
Figure 17-11b Accommodation.
b
For Distant Vision: Ciliary Muscle Relaxed, Lens Flattened
Lens flattened
Ciliary muscle
relaxed
© 2015 Pearson Education, Inc.
17-3 The Eye
•  Light Refraction of Lens
•  Image reversal
•  Visual acuity
•  Clarity of vision
•  “Normal” rating is 20/20
© 2015 Pearson Education, Inc.
Figure 17-12a Image Formation.
a
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Light from a point at the top
of an object is focused on
the lower retinal surface.
Figure 17-12b Image Formation.
b
© 2015 Pearson Education, Inc.
Light from a point at the
bottom of an object is
focused on the upper
retinal surface.
Figure 17-12c Image Formation.
Optic
nerve
c
© 2015 Pearson Education, Inc.
Light rays projected from a vertical
object show why the image arrives
upside down. (Note that the image
is also reversed.)
Figure 17-12d Image Formation.
Optic
nerve
d
© 2015 Pearson Education, Inc.
Light rays projected from a horizontal
object show why the image arrives with
a left and right reversal. The image also
arrives upside down. (As noted in the
text, these representations are not
drawn to scale.)
Figure 17-13 Refractive Problems (Part 1 of 5).
The eye has a fixed
focal distance and
focuses by varying the
shape of the lens.
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A camera lens has a
fixed size and shape
and focuses by varying
the distance to the film.
Figure 17-13 Refractive Problems (Part 2 of 5).
Emmetropia (normal vision)
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Figure 17-13 Refractive Problems (Part 3 of 5).
Myopia (nearsightedness)
If the eyeball is too deep or the resting
curvature of the lens is too great, the
image of a distant object is projected
in front of the retina. Myopic people
see distant objects as blurry and out
of focus. Vision at close range will be
normal because the lens is able to
round as needed to focus the image
on the retina.
Myopic
corrected with
a diverging,
concave
lens
© 2015 Pearson Education, Inc.
Diverging
lens
Figure 17-13 Refractive Problems (Part 4 of 5).
Hyperopia (farsightedness)
If the eyeball is too shallow or the lens
is too flat, hyperopia results. The
ciliary muscle must contract to focus
even a distant object on the retina.
And at close range the lens cannot
provide enough refraction to focus an
image on the retina. Older people
become farsighted as their lenses lose
elasticity, a form of hyperopia called
presbyopia (presbys, old man).
Hyperopia
corrected with
a converging,
convex
lens
© 2015 Pearson Education, Inc.
Converging
lens
Figure 17-13 Refractive Problems (Part 5 of 5).
Surgical Correction
Variable success at correcting myopia and
hyperopia has been achieved by surgery that
reshapes the cornea. In photorefractive
keratectomy (PRK) a computer-guided
laser shapes the cornea to exact specifications.
The entire procedure can be done in less than
a minute. A variation on PRK is called LASIK
(Laser-Assisted in-Situ Keratomileusis). In this
procedure the interior layers of the cornea are
reshaped and then recovered by the flap of original
outer corneal epithelium. Roughly 70 percent of LASIK
patients achieve normal vision, and LASIK has become the most
common form of refractive surgery.
Even after surgery, many patients still need reading glasses,
and both immediate and long-term visual problems can occur.
© 2015 Pearson Education, Inc.
17-4 Visual Physiology
•  Visual Physiology
•  Rods
•  Respond to almost any photon, regardless of
energy content
•  Cones
•  Have characteristic ranges of sensitivity
© 2015 Pearson Education, Inc.
17-4 Visual Physiology
•  Anatomy of Rods and Cones
•  Outer segment with membranous discs
•  Inner segment
•  Narrow stalk connects outer segment to inner
segment
© 2015 Pearson Education, Inc.
17-4 Visual Physiology
•  Anatomy of Rods and Cones
•  Visual pigments
•  Is where light absorption occurs
•  Derivatives of rhodopsin (opsin plus retinal)
•  Retinal synthesized from vitamin A
© 2015 Pearson Education, Inc.
Figure 17-14a Structure of Rods, Cones, and Rhodopsin Molecule.
Pigment Epithelium
In a cone, the discs are infoldings of
the plasma membrane, and the outer
segment tapers to a blunt point.
The pigment epithelium
absorbs photons that are
not absorbed by visual
pigments. It also
phagocytizes old discs
shed from the tip of the
outer segment.
In a rod, each disc is an independent
entity, and the outer segment forms
an elongated cylinder.
Melanin granules
Outer Segment
The outer segment of a
photoreceptor contains
flattened membranous
plates, or discs, that
contain the visual pigments.
Inner Segment
Discs
Connecting
stalks
Mitochondria
The inner segment contains
the photoreceptor’s major
organelles and is responsible
for all cell functions other
than photoreception. It also
releases neurotransmitters.
Golgi
apparatus
Nuclei
Cone
Rods
Each photoreceptor
synapses with a bipolar cell.
Bipolar cell
LIGHT
a Structure of rods and cones
© 2015 Pearson Education, Inc.
Figure 17-14b Structure of Rods, Cones, and Rhodopsin Molecule.
In a rod, each disc is an independent
entity, and the outer segment forms
an elongated cylinder.
Rhodopsin
molecule
Retinal
Opsin
b Structure of rhodospin molecule
© 2015 Pearson Education, Inc.
17-4 Visual Physiology
•  Color Vision
•  Integration of information from red, green, and
blue cones
•  Color blindness
•  Inability to detect certain colors
© 2015 Pearson Education, Inc.
Figure 17-15 Cone Types and Sensitivity to Color.
Light absorption
(percent of maximum)
100
Blue
cones
75
Rods
Red
Green cones
cones
50
25
0
W AV E L E N G T H ( n m )
400
450
500
550
600
650
Violet
Blue
Green
Yellow Orange
© 2015 Pearson Education, Inc.
700
Red
Figure 17-18 A Standard Test for Color Vision.
© 2015 Pearson Education, Inc.
17-4 Visual Physiology
•  Photoreception
•  Photon strikes retinal portion of rhodopsin
molecule embedded in membrane of disc
•  Opsin is activated
•  Bound retinal molecule has two possible
configurations
•  11-cis form
•  11-trans form
© 2015 Pearson Education, Inc.
Figure 17-16 Photoreception (Part 5 of 8).
1
Opsin activation occurs
The bound retinal molecule
has two possible
configurations: the 11-cis
form and the 11-trans form.
Photon
Rhodopsin
11-cis
retinal
11-trans
retinal
Opsin
Normally, the molecule is in
the curved 11-cis form; on
absorbing light it changes
to the more linear 11-trans
form. This change activates
the opsin molecule.
© 2015 Pearson Education, Inc.
Figure 17-16 Photoreception (Part 6 of 8).
2
Opsin activates
transducin, which
in turn activates
phosphodiesterase (PDE)
Transducin is a G
protein—a membranebound enzyme complex
Na+
PDE
Transducin
Disc
membrane
In this case, opsin activates
transducin, and transducin
in turn activates
phosphodiesterase (PDE).
© 2015 Pearson Education, Inc.
Figure 17-16 Photoreception (Part 7 of 8).
3
Cyclic GMP levels
decline and gated
sodium channels close
Phosphodiesterase is an
enzyme that breaks
down cGMP.
Na+
GMP
cGMP
The removal of cGMP from
the gated sodium channels
results in their inactivation.
The rate of Na+ entry into the
cytoplasm then decreases.
© 2015 Pearson Education, Inc.
Figure 17-16 Photoreception (Part 8 of 8).
IN LIGHT
ACTIVE STATE
−70 mV
4
Dark current is
reduced and rate of
neurotransmitter
release declines
The reduction in the rate
of Na+ entry reduces the
dark current. At the same
time, active transport
continues to export Na+
from the cytoplasm.
When the sodium
channels close, the
membrane potential
drops toward –70 mV. As the
plasma membrane
hyperpolarizes, the rate
of neurotransmitter
release decreases. This
decrease signals the
adjacent bipolar cell that
the photoreceptor has
absorbed a photon. After
absorbing a photon, retinal
does not spontaneously
revert to
he 11-cis form. Instead,
the entire rhodopsin
molecule must be broken
down into retinal and
opsin, in a process called
bleaching. It is then
reassembled.
© 2015 Pearson Education, Inc.
Na+
17-4 Visual Physiology
•  Recovery after Stimulation
•  Bleaching
•  Rhodopsin molecule breaks down into retinal and
opsin
•  Night blindness
•  Results from deficiency of vitamin A
© 2015 Pearson Education, Inc.
17-4 Visual Physiology
•  Light and Dark Adaptation
•  Dark
•  Most visual pigments are fully receptive to
stimulation
•  Light
•  Pupil constricts
•  Bleaching of visual pigments occurs
© 2015 Pearson Education, Inc.
17-4 Visual Physiology
•  The Visual Pathways
•  Begin at photoreceptors
•  End at visual cortex of cerebral hemispheres
•  Message crosses two synapses before it heads
toward brain
•  Photoreceptor to bipolar cell
•  Bipolar cell to ganglion cell
© 2015 Pearson Education, Inc.
17-4 Visual Physiology
•  Ganglion Cells
•  Monitor a specific portion of a field of vision
•  M Cells
•  Are ganglion cells that monitor rods
•  Are relatively large
•  Provide information about:
•  General form of object
•  Motion
•  Shadows in dim lighting
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17-4 Visual Physiology
•  Ganglion Cells
•  P cells
•  Are ganglion cells that monitor cones
•  Are smaller, more numerous
•  Provide information about edges, fine detail, and
color
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17-4 Visual Physiology
•  Ganglion Cells
•  On-center neurons
•  Are excited by light arriving in center of their
sensory field
•  Are inhibited when light strikes edges of their
receptive field
•  Off-center neurons
•  Inhibited by light in central zone
•  Stimulated by illumination at edges
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17-4 Visual Physiology
•  Central Processing of Visual Information
•  Axons from ganglion cells converge on optic disc
•  Penetrate wall of eye
•  Proceed toward diencephalon as optic nerve (II)
•  Two optic nerves (one for each eye) reach
diencephalon at optic chiasm
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17-4 Visual Physiology
•  Visual Data
•  From combined field of vision arrive at visual
cortex of opposite occipital lobe
•  Left half arrive at right occipital lobe
•  Right half arrive at left occipital lobe
•  Optic radiation
•  Bundle of projection fibers linking lateral
geniculates with visual cortex
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17-4 Visual Physiology
•  The Field of Vision
•  Depth perception
•  Obtained by comparing relative positions of objects
between left-eye and right-eye images
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17-4 Visual Physiology
•  The Brain Stem and Visual Processing
•  Circadian rhythm
•  Is tied to day–night cycle
•  Affects other metabolic processes
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Figure 17-20 The Visual Pathways.
Combined Visual Field
Left side
Left eye
only
Right side
Binocular vision
Right eye
only
The Visual
Pathway
Photoreceptors
in retina
Retina
Optic disc
Optic nerve
(II)
Optic chiasm
Optic tract
Lateral
geniculate
nucleus
Diencephalon
and
brain stem
Suprachiasmatic
nucleus
Projection fibers
(optic radiation)
Visual cortex
of cerebral
hemispheres
Left cerebral
hemisphere
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Superior
colliculus
Right cerebral
hemisphere
17-5 The Ear
•  The External Ear
•  Auricle
•  Surrounds entrance to external acoustic meatus
•  Protects opening of canal
•  Provides directional sensitivity
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17-5 The Ear
•  The External Ear
•  External acoustic meatus
•  Ends at tympanic membrane (eardrum)
•  Tympanic membrane
•  Is a thin, semitransparent sheet
•  Separates external ear from middle ear
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Figure 17-21 The Anatomy of the Ear.
Middle Ear
External Ear
Elastic cartilages
Internal Ear
Auditory ossicles
Oval
window
Semicircular canals
Petrous part of
temporal bone
Auricle
Facial nerve (VII)
Vestibulocochlear
nerve (VIII)
Bony labyrinth
of internal ear
Cochlea
Tympanic
cavity
Auditory tube
To
nasopharynx
External acoustic
meatus
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Tympanic
membrane
Round
window
Vestibule
17-5 The Ear
•  The External Ear
•  Ceruminous glands
•  Integumentary glands along external acoustic
meatus
•  Secrete waxy material (cerumen)
•  Keeps foreign objects out of tympanic membrane
•  Slows growth of microorganisms in external acoustic
meatus
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Figure 17-22a The Middle Ear.
Auditory Ossicles
Malleus Incus Stapes
Temporal bone
(petrous part)
Oval window
Stabilizing
ligaments
Muscles of
the Middle Ear
Branch of facial
nerve VII (cut)
Tensor tympani
muscle
External
acoustic meatus
Stapedius muscle
Tympanic cavity
(middle ear)
Round window
Auditory tube
Tympanic
membrane
a
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The structures of the middle ear
Figure 17-22b The Middle Ear.
Malleus
attached to
tympanic
membrane
Malleus
Tendon of tensor
tympani muscle
Incus
Base of stapes
at oval window
Stapes
Stapedius muscle
Inner surface of
tympanic membrane
b The tympanic membrane and auditory ossicles
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Figure 17-22c The Middle Ear.
Incus
Malleus
Points of
attachment
to tympanic
membrane
Stapes
Base of stapes
c The isolated auditory ossicles
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Figure 17-23b The Internal Ear.
KEY
Membranous
labyrinth
Semicircular ducts
Bony labyrinth
Anterior
Lateral
Vestibule
Posterior
Cristae within ampullae
Maculae
Endolymphatic sac
Semicircular
canal
Cochlea
Utricle
Saccule
Vestibular duct
Cochlear duct
Tympanic
duct
Spiral
organ
b The bony and membranous labyrinths. Areas of
the membranous labyrinth containing sensory
receptors (cristae, maculae, and spiral organ) are
shown in purple.
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17-5 The Ear
•  Equilibrium
•  Sensations provided by receptors of vestibular
complex
•  Hair cells
•  Basic receptors of inner ear
•  Provide information about direction and strength of
mechanical stimuli
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17-5 The Ear
•  The Semicircular Ducts
•  Are continuous with utricle
•  Each duct contains:
•  Ampulla with gelatinous cupula
•  Associated sensory receptors
•  Stereocilia – resemble long microvilli
•  Are on surface of hair cell
•  Kinocilium – single large cilium
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Figure 17-24a The Semicircular Ducts.
Vestibular branch (N VIII)
Semicircular ducts
Anterior
Cochlea
Ampulla
Posterior
Endolymphatic sac
Lateral
Endolymphatic duct
Utricle
Maculae
Saccule
a An anterior view of the right
semicircular ducts, the utricle,
and the saccule, showing the
locations of sensory receptors.
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Figure 17-24b The Semicircular Ducts.
Ampulla
filled with
endolymph
Cupula
Hair cells
Crista ampullaris
Supporting cells
Sensory nerve
b A cross section through the ampulla of a semicircular duct.
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Figure 17-24c The Semicircular Ducts.
Direction of
duct rotation
Direction of relative
endolymph movement
Direction of
duct rotation
Semicircular duct
Cupula
At rest
c Endolymph movement along the length of the duct
moves the cupula and stimulates the hair cells.
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Figure 17-24d The Semicircular Ducts.
Displacement in
this direction
stimulates hair cell
Displacement in
this direction
inhibits hair cell
Gelatinous
material
Kinocilium
Stereocilia
Hair cell
Sensory nerve
ending
Supporting cell
d
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A representative hair cell (receptor) from the
vestibular complex. Bending the sterocilia toward
the kinocilium depolarizes the cell and stimulates
the sensory neuron. Displacement in the opposite
direction inhibits the sensory neuron.
17-5 The Ear
•  The Utricle and Saccule
•  Provide equilibrium sensations
•  Are connected with the endolymphatic duct,
which ends in endolymphatic sac
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17-5 The Ear
•  The Utricle and Saccule
•  Maculae
•  Oval structures where hair cells cluster
•  Statoconia
•  Densely packed calcium carbonate crystals on
surface of gelatinous mass
•  Otolith (ear stone) = gelatinous matrix and
statoconia
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Figure 17-25ab The Saccule and Utricle.
Endolymphatic sac
Endolymphatic duct
Utricle
Saccule
a The location of
the maculae
Otoliths
Gelatinous layer
forming otolithic
membrane
Hair cells
Nerve fibers
b The structure of an individual macula
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Figure 17-25c The Saccule and Utricle.
1
Head in normal, upright position
Gravity
2
Head tilted posteriorly
Receptor
output
increases
c
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Gravity
Otolith
moves
“downhill,”
distorting hair
cell processes
A diagrammatic view of utricular macular function
when the head is held normally 1 and then tilted
back 2
17-5 The Ear
•  Four Functions of Vestibular Nuclei
1.  Integrate sensory information about balance and
equilibrium from both sides of head
2.  Relay information from vestibular complex to
cerebellum
3.  Relay information from vestibular complex to
cerebral cortex
•  Provide conscious sense of head position and
movement
4.  Send commands to motor nuclei in brain stem
and spinal cord
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Figure 17-26 Pathways for Equilibrium Sensations.
To superior colliculus and
relay to cerebral cortex
Red nucleus
N III
Semicircular
canals
Vestibular
ganglion
N IV
Vestibular
branch
Vestibular
nucleus
N VI
To
cerebellum
Vestibule
Cochlear
branch
N XI
Vestibulocochlear
nerve (VIII)
Vestibulospinal
tracts
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17-5 The Ear
•  Eye, Head, and Neck Movements
•  Reflexive motor commands
•  From vestibular nuclei
•  Distributed to motor nuclei for cranial nerves
•  Peripheral Muscle Tone, Head, and Neck
Movements
•  Instructions descend in vestibulospinal tracts of
spinal cord
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17-5 The Ear
•  Eye Movements
•  Sensations of motion directed by superior colliculi
of the midbrain
•  Attempt to keep focus on specific point
•  If spinning rapidly, eye jumps from point to point
•  Nystagmus
•  Have trouble controlling eye movements
•  Caused by damage to brain stem or inner ear
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17-5 The Ear
•  Hearing
•  Cochlear duct receptors
•  Provide sense of hearing
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Figure 17-27a The Cochlea.
Round window
Stapes at
oval window
Scala
vestibuli
Cochlear
duct
Scala
tympani
Semicircular
canals
KEY
Cochlear
branch
Vestibular
branch
Vestibulocochlear
nerve (VIII)
a The structure of the cochlea
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From oval window
to tip of spiral
From tip of spiral
to round window
Figure 17-27b The Cochlea.
Temporal bone
(petrous part)
Vestibular
membrane
Scala vestibuli
(contains perilymph)
Tectorial
membrane
Cochlear duct
(contains endolymph)
Basilar
membrane
Spiral organ
From oval
window
Spiral ganglion
Scala tympani
(contains perilymph)
To round
window
Cochlear nerve
Vestibulocochlear nerve (VIII)
b Diagrammatic and sectional views of the cochlear spiral
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17-5 The Ear
•  Hearing
•  Auditory ossicles
•  Convert pressure fluctuation in air into much
greater pressure fluctuations in perilymph of
cochlea
•  Frequency of sound
•  Determined by which part of cochlear duct is
stimulated
•  Intensity (volume)
•  Determined by number of hair cells stimulated
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17-5 The Ear
•  Hearing
•  Cochlear duct receptors
•  Basilar membrane
•  Separates cochlear duct from tympanic duct
•  Hair cells lack kinocilia
•  Stereocilia in contact with overlying tectorial
membrane
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Figure 17-28a The Spiral Organ.
Bony cochlear wall
Scala vestibuli
Vestibular membrane
Spiral
ganglion
Cochlear duct
Tectorial membrane
Basilar membrane
Scala tympani
Spiral organ
a A three-dimensional section of the
cochlea, showing the compartments,
tectorial membrane, and spiral organ
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Cochlear branch
of N VIII
Figure 17-28b The Spiral Organ (Part 1 of 2).
Tectorial membrane
Outer
hair cell
Basilar
membrane
Inner
hair cell
Nerve
fibers
b Diagrammatic and sectional views of the receptor
hair cell complex of the spiral organ
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17-5 The Ear
•  An Introduction to Sound
•  Pressure waves
•  Consist of regions where air molecules are
crowded together
•  Adjacent zone where molecules are farther apart
•  Sine waves
•  S-shaped curves
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17-5 The Ear
•  Pressure Wave
•  Wavelength
•  Distance between two adjacent wave troughs
•  Frequency
•  Number of waves that pass fixed reference point at
given time
•  Physicists use term cycles instead of waves
•  Hertz (Hz) number of cycles per second (cps)
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17-5 The Ear
•  Pressure Wave
•  Pitch
•  Our sensory response to frequency
•  Amplitude
•  Intensity of sound wave
•  Sound energy is reported in decibels
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Figure 17-29a The Nature of Sound.
Wavelength
Tympanic
membrane
Tuning
fork
Air
molecules
a Sound waves (here, generated by a tuning fork)
travel through the air as pressure waves.
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Sound energy arriving at
tympanic membrane
Figure 17-29b The Nature of Sound.
1 wavelength
Amplitude
Time (sec)
b
A graph showing the sound energy arriving at the
tympanic membrane. The distance between wave
peaks is the wavelength. The number of waves
arriving each second is the frequency, which we
perceive as pitch. Frequencies are reported in
cycles per second (cps), or hertz (Hz). The amount
of energy carried by the wave is its amplitude.
The greater the amplitude, the louder the sound.
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Figure 17-30 Sound and Hearing (Part 1 of 2).
External
acoustic
meatus
Malleus
Incus
Stapes
Oval
window
3
Movement
of sound
waves
2
1
Round
window
Tympanic
membrane
1
Sound waves
arrive at
tympanic
membrane.
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3
2
Movement of
the tympanic
membrane causes
displacement
of the auditory
ossicles
Movement of
the stapes at
the oval window
establishes
pressure waves
in the perilymph
of the scala
vestibuli.
Figure 17-30 Sound and Hearing (Part 2 of 2).
Cochlear branch
of cranial nerve VIII
6
Scala vestibuli
(contains perilymph)
Vestibular membrane
Cochlear duct
(contains endolymph)
Basilar membrane
4
Scala tympani
(contains perilymph)
5
4
The pressure
waves distort
the basilar
membrane on
their way to the
round window
of the scala
tympani.
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6
5
Vibration of
the basilar
membrane
causes hair cells
to vibrate against
the tectorial
membrane.
Information about
the region and
the intensity of
stimulation is
relayed to the CNS
over the cochlear
branch of cranial
nerve VIII.
17-5 The Ear
•  Auditory Pathways
•  Cochlear branch
•  Formed by afferent fibers of spiral ganglion
neurons
•  Enters medulla oblongata
•  Synapses at dorsal and ventral cochlear nuclei
•  Information crosses to opposite side of brain
•  Ascends to inferior colliculus of midbrain
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17-5 The Ear
•  Auditory Pathways
•  Ascending auditory sensations
•  Synapse in medial geniculate nucleus of thalamus
•  Projection fibers deliver information to auditory
cortex of temporal lobe
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Figure 17-32 Pathways for Auditory Sensations (Part 1 of 2).
1
Stimulation of hair cells at
a specific location along
the basilar membrane
activates sensory neurons.
Cochlea
Low-frequency
sounds
High-frequency
sounds
Vestibular
branch
2
Sensory neurons carry the
sound information in the
cochlear branch of the
vestibulocochlear nerve
(VIII) to the cochlear
nucleus on that side.
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KEY
Vestibulocochlear
nerve (VIII)
Primary pathway
Secondary pathway
Motor output
17-5 The Ear
•  Hearing Range
•  From softest to loudest represents trillionfold
increase in power
•  Never use full potential
•  Young children have greatest range
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Table 17-1 Intensity of Representative Sounds.
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17-5 The Ear
•  Effects of Aging on the Ear
•  With age, damage accumulates
•  Tympanic membrane gets less flexible
•  Articulations between ossicles stiffen
•  Round window may begin to ossify
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