Aand PCH 15 Lecture Pearson

15
The Special Senses
Lecture Presentation by
Lori Garrett
© 2018 Pearson Education, Inc.

Note to the Instructor:
For the third edition of Visual Anatomy & Physiology, we have updated our PowerPoints to
fully integrate text and art. The pedagogy now more closely matches that of the textbook.
The goal of this revised formatting is to help your students learn from the art more
effectively. However, you will notice that the labels on the embedded PowerPoint art are
not editable. You can easily import editable art by doing the following:
Copying slides from one slide set into another

You can easily copy the Label Edit art into the Lecture Presentations by using either the PowerPoint
Slide Finder dialog box or Slide Sorter view. Using the Slide Finder dialog box allows you to explicitly
retain the source formatting of the slides you insert.
Using the Slide Finder dialog box in PowerPoint:
1. Open the original slide set in PowerPoint.
2. On the Slides tab in Normal view, click the slide thumbnail that you want the copied slides to
follow.
3. On the toolbar at the top of the window, click the drop down arrow on the New Slide tab. Select
Reuse Slides.
4. Click Browse to look for the file; in the Browse dialog box, select the file, and then click Open.
5. If you want the new slides to keep their current formatting, in the Slide Finder dialog box, select
the Keep source formatting checkbox. When this checkbox is cleared, the copied slides assume
the formatting of the slide they are inserted after.
6. To insert selected slides: Click the slides you want to insert. Slides will place immediately after the
slide you have selected in the Slides tab in Normal view.
Section 1: Olfaction and Gustation
Learning Outcomes
15.1 Explain the roles of generator potentials and
depolarization in sensory neurons and receptor
cells.
15.2 Describe the sensory organs of smell, trace the
olfactory pathways to their destinations in the
cerebrum, and explain how olfactory perception
occurs.
15.3 Describe the sensory organs of gustation.
15.4 Describe gustatory reception, briefly describe
the physiological processes involved in taste,
and trace the gustatory pathway.
Module 15.1: A generator potential is a
depolarization of the membrane
 Five special senses
1. Olfaction (smell)
2. Gustation (taste)
3. Vision
4. Equilibrium (balance)
5. Hearing
 All originate with one of two types of sensory
receptor cells
1. Dendrites of specialized neurons
– Example: olfactory receptors
2. Specialized cells that synapse with sensory neurons
 Depolarization of sensory neuron = a generator
potential
Module 15.1: Generator potential
1. Olfactory receptors are dendrites of specialized

neurons
 Dissolved odorants bind to olfactory receptors
 Triggers depolarization = generator potential
 With strong enough stimulus, generator potential
triggers action potentials that go to CNS

2. Receptors for taste, vision, equilibrium, hearing

are specialized cells with inexcitable
membranes
 Synapse with sensory neurons
 Stimulation—triggers graded depolarization

 Graded depolarization of the receptor cell triggers

neurotransmitter release
 Neurotransmitter depolarizes sensory neurons,
causing generator potential that can trigger action
potential
 Action potentials are propagated to CNS

Module 15.1: Review
A. Where do the special senses originate?

B. What is a generator potential?
C. Compare olfactory receptors with receptors for
the other special senses.
Learning Outcome: Explain the roles of generator

potentials and depolarization in sensory neurons
and receptor cells.

Module 15.2: Olfaction involves specialized
chemoreceptive neurons and delivers
sensations directly to the cerebrum
Olfaction = sense of smell
 Paired olfactory organs in nasal cavity, each side
of nasal septum
• Contain olfactory receptor cells
• Distributed along cribriform plate, superior portion of
the perpendicular plate, superior
nasal conchae
 Olfactory organs have two layers
1. Olfactory epithelium
2. Lamina propria

Module 15.2: Olfaction
Olfactory epithelium
 Olfactory receptor cells (modified neurons)
• Each forms knob with up to 20 cilia-shaped dendrites
projecting past epithelial surface into mucus
• 10–20 million receptors in a 5-cm2 area
 Supporting cells
 Basal cells (stem cells)—constantly produce new
receptor cells
• One of the few examples of neuronal replacement

Lamina propria
 Areolar tissue, blood vessels, nerves
 Olfactory glands—secretions absorb water; form
thick, pigmented mucus

An olfactory organ, showing the olfactory
epithelium and lamina propria

Odorants = dissolved chemicals that stimulate

olfactory neurons
 Bind membrane receptors (odorant-binding proteins)
on dendrites of olfactory receptor cells
 Generally small organic molecules
 As few as four odorant molecules can activate
receptor cell

Olfactory reception process

1. Odorant binds to receptor protein; activates
adenylate cyclase (enzyme that converts ATP to
cyclic AMP)
2. cAMP opens sodium channels in plasma
membrane; starts depolarization
3. If enough depolarization occurs, action potential is
triggered, and information is relayed to CNS

The process of olfaction



Olfactory pathway to the cerebrum

1. Chemicals in air bind odorant-binding proteins on
olfactory cell membranes; stimulate sensory
neurons in olfactory organ
2. Axons leave olfactory epithelium in bundles
passing through cribriform plate (ethmoid)
3. First synapse occurs in olfactory bulb
immediately superior to cribriform plate
4. Axons leaving olfactory bulb travel through
olfactory tract to olfactory cortex, hypothalamus,
and limbic system
• Distribution to limbic system (emotions) and
hypothalamus (ANS) explains why smells can produce
profound emotional and behavioral responses
The olfactory pathway





Module 15.2: Review
A. Describe olfaction.
B. Trace the olfactory pathway, beginning at the
olfactory epithelium.
C. Describe the events leading up to the
depolarization and generation of an action
potential by an olfactory receptor cell.
Learning Outcome: Describe the sensory organs of

smell, trace the olfactory pathways to their
destinations in the cerebrum, and explain how
olfactory perception occurs.

Module 15.3: Gustation involves epithelial
chemoreceptor cells located in taste buds
Gustation = taste
 Taste receptors (gustatory receptor cells)
• Most important receptors on superior surface of
tongue
• Also in adjacent parts of pharynx, larynx, epiglottis
– Numbers decrease with age

Module 15.3: Gustation involves epithelial
chemoreceptor cells located in taste buds
Gustation = taste (continued)

 Lingual papillae = epithelial projections on tongue
surface
• Contain taste buds = sensory structures with taste
receptors that respond to various chemical stimuli,
and specialized epithelial cells.
• Four types of lingual papillae: vallate papillae, foliate
papillae, fungiform papillae, and filiform papillae

Module 15.3: Gustation

 Vallate (vallum, wall) papillae
• Relatively large—each contains up to 100 taste buds
• Surrounded by deep epithelial folds
• Form “V” at back of tongue


 Foliate papillae—lateral margins of posterior region
of tongue
 Fungiform (fungus, mushroom) papillae—about
five taste buds each


 Filiform (filum, thread) papillae—provide friction to
help tongue move objects around mouth
• Anterior two-third of superior surface of tongue
• NO taste buds


Four primary taste sensations: sweet, salty,

sour, bitter
 No difference in taste
bud structure
 Taste buds in all portions
of tongue provide all four
sensations
 Umami = pleasant,
savory taste characteristic
of broths, cheese
• Binds receptors for
amino acids, small peptides, nucleotides

Four primary taste sensations: sweet, salty,

sour, bitter (continued)
 Water receptors
• Concentrated in pharynx
• Output goes to hypothalamus;
affects water balance
and regulation of
blood volume
• Prevent overingesting
H2O

Taste buds
 Recessed into epithelium
 Gustatory receptor cell
(taste receptor cell)
• Extends slender microvilli (taste
hairs) into surrounding fluids
through a taste pore
• Typically lives about 10 days
before it is replaced
• About 40–100 gustatory cells in each taste bud
 Basal cells = stem cells that divide and mature to
produce transitional cells that mature into new
gustatory cells

Taste receptor sensitivity

 Threshold varies for the four primary sensations
 More sensitive to unpleasant stimuli
• 100,000 times more sensitive to bitter; 1000 times
more sensitive to sour (acids) than to sweet or salty
• Survival value—acids burn tissues; many toxins are
bitter
• Overall sensitivity declines
with age—leads to changing
taste sensations with age

Module 15.3: Review
A. Define gustation.
B. Describe filiform papillae.
C. Which taste receptors offer a survival
advantage when tasting something for the first
time?
Learning Outcome: Describe the sensory organs of

gustation.

Module 15.4: Gustatory reception relies on
membrane receptors and ion channels, and
sensations are carried by facial,
glossopharyngeal, and vagus nerves
Gustatory reception stimulated by dissolved

chemicals (like smell)
 Chemicals contacting taste hairs may:
• Diffuse through plasma membrane leak channels
• Bind to receptor proteins of gustatory receptor cell
 ~ 90 percent of gustatory receptor cells respond to
at least two taste stimuli
 Different tastes involve different receptor
mechanisms

Module 15.4: Gustatory reception
Two types of gustatory reception

1. Salt and sour receptors
• Sodium ions (salt) or hydrogen ions
(sour) diffuse through Na+ leak
channels
• Membrane/cell depolarizes;
neurotransmitter is released
 Neurotransmitters are released at
synapse with sensory neurons
 Depolarization of sensory neurons
leads to generator potential, which
can produce action potentials along
gustatory pathway to CNS
2. Sweet, bitter, and umami receptors:

• Binding to these receptors activates
G-protein complexes called
gustducins (protein complexes that
use second messengers to produce
effects)
• Activated second messenger causes
neurotransmitter release
 Neurotransmitters are released at
synapse with sensory neurons
 Depolarization of sensory neurons
leads to generator potential, which can produce
action potentials along gustatory pathway to CNS
Different tastes involve different receptor
mechanisms

Gustatory pathway—gustatory receptor cells to

cerebral cortex
1. Gustatory receptor cells respond to stimulation
2. Information is relayed on cranial nerves VII, IX,
and X
3. Sensory afferents synapse in solitary nucleus of
medulla oblongata
4. Postsynaptic neurons cross over; enter medial
lemniscus of medulla oblongata
5. Synapse in thalamus, then go to gustatory cortex
in insula for conscious perception

Gustatory pathway (continued)

1. Gustatory receptor cells
bind dissolved
chemicals
• Generator potential
occurs
• Triggers action potentials


2. Information is relayed
on cranial nerves
• Facial nerve (VII)—taste
buds on anterior two-
thirds of tongue, from
the tip to the vallate
papillae
• Glossopharyngeal nerve
(IX)—vallate papillae,
posterior one-third of tongue
• Vagus nerve (X)—epiglottis


3. Sensory afferents
synapse in solitary
nucleus of medulla
oblongata


4. Axons of postsynaptic
neurons cross over;
enter medial
lemniscus of
medulla oblongata


5. Synapse in thalamus;
then information is
projected to appropriate
portions of gustatory
cortex of the insula

Taste = conscious perception produced by

processing at the primary somatosensory cortex
 Information from taste buds is integrated with other
sensory data
• Texture of food
• Taste-related sensations (“peppery” or “burning
hot”)—carried by trigeminal nerve (V)
 Taste receptors adapt slowly, but central adaptation
reduces sensitivity to a new taste quickly
 Level of olfactory stimulation plays important role
• Several thousand times more sensitive to “tastes”
when sense of smell is functioning

Module 15.4 Review
A. What are gustducins?

B. Identify the cranial nerves that carry gustatory
information.
C. Trace the gustatory pathway from the taste
receptors to the cerebral cortex.
Learning Outcome: Describe gustatory reception,

briefly describe the physiological processes
involved in taste, and trace the gustatory pathway.

Section 2: Vision
Learning Outcomes
15.5 Outline the embryonic development of the eye.
15.6 Identify the accessory structures of the eye, and
explain their functions.
15.7 Describe the layers of the eye wall and the
anterior and posterior cavities of the eye.
15.8 Explain how light is directed to the fovea
centralis of the retina.
15.9 Describe the process by which images are
focused on the retina.

Section 2: Vision
Learning Outcomes (continued)

15.10 Describe the structure and function of the retina’s
layers of cells, and explain the distribution of rods
and cones and their relation to visual acuity.
15.11 Describe the structure of the photoreceptors and
how we are able to distinguish colors.
15.12 Explain photoreception and how visual pigments
are activated.

Section 2: Vision

15.13 Explain how the visual pathways distribute
information to their destinations in the brain.
15.14 Clinical Module: Describe various refraction
problems associated with the cornea, lens, or
shape of the eye.

Module 15.5: The eyes form early in embryonic
development
 Eyes are our most complex

sense organs
 Much processing occurs in eye
before relayed to CNS
 Resemble detached CNS
nuclei that process visual
stimuli
Eye development
1. Begins as pair of bulges = optic vesicles
• Each contains cavity continuous with neural tube
• Form in prosencephalon walls

Module 15.5: Eye development
Eye development (continued)

2. Lateral bulges indent; form
optic cups—connected to
diencephalon
• Overlying epidermis forms
pocket; pinches off to form
the lens
• Inner and outer layers form
retina
– Outer ependymal cells become photoreceptors
– Inner ependymal cells become pigment cells
– Nervous tissue of outer layer forms neurons, ganglion
cells, specialized glial cells

Module 15.5: Eye development
Eye development (continued)

3. Embryonic cells around optic
cup form supporting layers of
connective tissue
• Isolate nervous tissue
• Interior chambers develop;
filled with fluid

Module 15.5: Review
A. What are the first structures that form during

eye development?
B. Which structures develop into the retina?
C. Which cells develop into the photoreceptors?
Learning Outcome: Outline the embryonic

development of the eye.

Module 15.6: Accessory structures of the eye
provide protection while allowing light to reach
the interior of the eye
Eyelashes = robust hairs that keep foreign matter
out of eyes
Eyelids (palpebrae) = continuation of skin; can
close to protect eye
 Blinking lubricates eye surface; clears dust/debris
 Medial angle of the eye = where two eyelids meet
medially

Module 15.6: Eye accessory structures
 Lateral angle of the eye = where two eyelids meet

laterally
 Palpebral fissure = gap between upper/lower
eyelids

 Cornea = transparent area on anterior surface

 Pupil = opening in center of iris; transmits light
 Lacrimal caruncle = small, reddish body at medial
angle
• Produces thick secretions that cause gritty deposits
sometimes appearing after night’s sleep

Conjunctiva = mucous membrane and epithelium

lining eyelids and covering anterior of eye
 Palpebral conjunctiva—inner surface of eyelids
 Bulbar conjunctiva—anterior eye surface;
continuous with cornea
 Fornix = pocket created at junction of conjunctiva
Tarsal glands = modified
sebaceous glands
 Inner margin of eyelids
 Secretions prevent eyelid
sticking

Conjunctivitis, or pinkeye
 Inflammation of the conjunctiva
 Redness due to dilation of blood vessels deep to
conjunctival epithelium
 From pathogenic infection or physical, allergic, or
chemical irritation of conjunctival surface

Lacrimal apparatus
 Lacrimal gland
• ~12–20 mm long
• Produces tears that lubricate, nourish, oxygenate,
clean cornea
• Contain lysozyme and
antibodies—attack
invading pathogens

Lacrimal apparatus (continued)

 Tear ducts—10–12 tubes; deliver tears from
lacrimal gland to space behind upper eyelid (forming
lacrimal “lake”)
 Lacrimal puncta—two small pores that drain the
lacrimal lake
 Lacrimal canaliculi = connect
lacrimal puncta to
lacrimal sac

Lacrimal apparatus (continued)

 Lacrimal sac = small chamber nestled in lacrimal
sulcus of orbit
 Nasolacrimal duct
• From inferior tip of lacrimal sac through nasolacrimal
canal
• Delivers tears to nasal cavity
• Empties into inferior
meatus (inferior/
lateral to inferior
nasal concha)

Module 15.6 Review
A. Identify the accessory structures of the eye.

B. Which layer of the eye would be the first
affected by inadequate tear production?
C. Describe conjunctivitis.
Learning Outcome: Identify the accessory

structures of the eye, and explain their functions.

Module 15.7: The hollow eyeball has a layered
wall and fluid-filled anterior and posterior
cavities
Three layers (tunics) of the eye
1. Outer fibrous layer—sclera, corneal limbus, and
cornea
2. Middle vascular layer—
iris, choroid, ciliary body
3. Inner layer—retina

Module 15.7: Eye layers and cavities
Three layers (tunics) of the eye (continued)

1. Fibrous layer
• Outermost layer of eye
• Consists of cornea (clear) and sclera (white)
– Joined at corneoscleral junction
• Functions
1. Supports and protects the eye
2. Attachment site for extrinsic eye muscles
3. Curvature of the cornea aids in the focusing process
(light first enters through cornea)


2. Vascular layer (uvea)—contains many blood
vessels, lymphatic vessels, and intrinsic (smooth)
muscles of eye
• Includes iris, ciliary body, and choroid
• Functions
1. Provides route for blood vessels/lymphatics to eye
tissues
2. Regulates amount of light entering eye (iris)
3. Secretes/reabsorbs aqueous humor (fluid) circulating
in eye chambers
4. Controls shape of lens (ciliary body); essential for
focusing


 Iris = colored part of eye
• Blood vessels, pigment cells (melanocytes)
• Two layers of smooth muscle—contraction changes
diameter of pupil to control amount of light entering
 Ciliary body = thickened region bulging into interior
of eye
• Ring of fibers connects ciliary body and lens
 Choroid = vascular layer underlying sclera; has
extensive capillary network supplying oxygen/
nutrients to neural layer


3. Inner layer, or retina = innermost layer of eye
• Outer pigmented layer absorbs light
• Thick, inner neural layer contains photoreceptors
(the cells sensitive to light)

The three layers of the eye and associated
structures

Eye cavities—separated by lens/ciliary body

 Anterior cavity extends from cornea to lens;
contains fluids called aqueous humor
• Two chambers
1. Anterior chamber—cornea to iris
2. Posterior chamber (iris to ciliary body and lens)
 Posterior cavity is filled with gelatinous vitreous
body
• Vitreous humor = fluid
part of vitreous body

The anterior cavity

 Ciliary body
1. Ciliary muscle—smooth muscle ring, projects inward
2. Ciliary processes—folds of epithelium covering
ciliary muscles
3. Ciliary zonule (suspensory ligaments)—holds lens
in position so light passes through it

The anterior cavity (continued)

 Aqueous humor
• Circulates within anterior cavity, passes through pupil
• Secreted by epithelial cells of ciliary processes
• Diffuses through vitreous body to retinal surface
(vitreous humor)

Aqueous humor (continued)

 Leaves eye at scleral venous sinus (canal of
Schlemm) = passageway around eye at level of
corneoscleral junction
 Flows into veins in sclera
 Rate of removal should keep pace with rate of
secretion

Aqueous humor (continued)

 Functions
1. Transports nutrients and wastes
2. Forms fluid cushion
3. Helps retain eye shape
4. Stabilizes position of the retina

Iris
 Body of iris is highly vascular, pigmented loose
connective tissue
• Anterior surface—incomplete layer of
fibroblasts/melanocytes
• Posterior surface—lined by pigmented epithelium of
neural layer

Iris (continued)
• Oro serrata—jagged edge of neural layer of retina
• Eye color—determined by genes that influence
density and distribution of melanocytes and density of
pigmented epithelium

Module 15.7: Review
A. Name the three layers of the eye.

B. What give eyes their characteristic color?
C. Where in the eye is aqueous humor located?
Learning Outcome: Describe the layers of the eye

wall and the anterior and posterior cavities of the
eye.

Module 15.8: The structures of the eye direct
light along a visual axis to the fovea centralis of
the retina
Cornea
 Allows light to enter eye—transparent and clear
 Dense matrix of multiple layers of collagen fibers
 Avascular; receives oxygen and nutrients from tears

Module 15.8: Eye structures
Lens
 Posterior to cornea; anchored by ciliary zonule of
ciliary body,
 Concentric layers of cells filled with transparent
crystallins = proteins responsible for clarity and
focusing power of lens
 Dense fibrous capsule
surrounds cells; blends
with ciliary zonule
 Primary function:
changes shape to
focus image on
photoreceptors
 Choroid = middle layer; blood vessels nourish all eye

structures
 Sclera (“white of the eye”)—dense fibrous connective
tissue with collagen and elastic fibers
• Stabilizes eye shape during movement
• Insertion for extrinsic eye muscles
 Optic nerve (II)—conveys
visual information to brain

 Ciliary body—supports lens, controls its shape;

tension in ciliary zonule resists tendency of lens to
ball up
 Retina—contains photoreceptors, pigment cells,
supporting cells, neurons

Pupil = opening in iris through which light passes

 Two pupillary muscles of iris regulate amount of light
entering
1. Dilator pupillae muscles—extend radially from
pupil
– Enlarge pupil; supplied by sympathetic nervous system

2. Sphincter pupillae muscles—encircle pupil like
a ring
– Make pupil smaller; supplied by parasympathetic
nervous system

Action of the dilator pupillae muscle

Action of the sphincter pupillae muscle

Visual axis =
imaginary line drawn
from center of object
you are looking at
through the center of
the cornea and lens to
retina

Retina—photoreceptors, pigment cells, supporting

cells, neurons
 Photoreceptors in inner neural portion; type/density
vary by area
• Macula—patch of retina with high density of
photoreceptors
• Fovea centralis—central
part of macula
– Has highest concentration
of photoreceptors
– Point of sharpest vision

Review of structures of the eye

Module 15.8: Review
A. Which eye structure does not contain blood

vessels?
B. What happens to the pupils when light intensity
decreases?
C. Light passing through the eye along the visual
axis strikes what part of the retina?
Learning Outcome: Explain how light is directed to

the fovea centralis of the retina.

Module 15.9: Focusing of light produces a
sharp image on the retina
Focusing process is a two-step process

1. Light is refracted (bent) when it passes from air
into cornea
• Bending occurs because of the change in density
• Amount of refraction at cornea is constant
2. More refraction when light passes from aqueous
humor into lens
• Bends light rays toward focal point—specific point on
retina

Module 15.9: Focusing on the retina
Focal distance of a lens

 Distance between center of lens and its focal point
 Determined by:
1. Distance from object to lens
2. Shape of lens
 Distance from lens to retina cannot change
• Focus by changing shape of the lens =
accommodation

Accommodation = change in lens shape to keep

focal distance constant and provide clear vision
 For close vision:
• Ciliary muscle contracts—ciliary body moves toward
lens, reduces tension in ciliary zonule
• Lens pulled into more spherical shape; increases
refraction
• Light from near
objects is focused
on retina

Accommodation (continued)
 For distant vision:
• Ciliary muscle relaxes—ciliary zonule pulls on lens
• Lens flattens; brings image of distant object into focus
on retina

Image formation
 Images are not single point, but rather consist of
large numbers of individual points (like pixels on
computer screen)—each focused on retina
 Image is inverted and reversed; brain
compensates—learned from experience

Near point of vision = inner limit of clear vision

 Determined by lens elasticity
 Increases with age as lens becomes less elastic
• In children 7–9 cm (3–4 in.)
• Young adult 15–20 cm (6–8 in.)
• Age 60, about 83 cm (33 in.)

Module 15.16: Review
A. Define focal point.

B. When the ciliary muscles are relaxed, are you
viewing something close up or something in the
distance?
C. Why does the near point of vision typically
increase with age?
Learning Outcome: Describe the process by which

images are focused on the retina.

Module 15.10: The neural layer of the retina
contains multiple layers of specialized
photoreceptors, neurons, and supporting cells
Retinal layers—pigmented and neural layers
 Pigmented layer of the retina—absorbs light that
passes through neural part to prevent light bouncing
back, producing visual “echoes”

Module 15.10: Retinal layers and cells
Retinal layers (continued)

 Neural layer of the retina—photoreceptors,
supporting cells, neurons
• Preliminary processing/integration of visual
information
• Photoreceptors located closest to pigmented layer

Retinal layers (continued)

 Ganglion cells—innermost layer
• Axons converge at optic disc—form optic nerve
(optic disc also called blind spot—lacks
photoreceptors)
• Blood vessels follow optic nerve, supply inner neural
layers cells


Photoreceptors of the retina

 Rods
• Highly sensitive; allow vision in very dim light
• No color discrimination—provide black-and-white
vision only
 Cones
• Color vision
• Sharper, clearer images
• Require more intense light
 Rods and cones synapse with bipolar cells; bipolar
cells synapse on ganglion cells

Photoreceptors of the retina

Horizontal and amacrine cells

 Facilitate/inhibit communication between
photoreceptors and ganglion cells
 Horizontal cells—where photoreceptors/bipolar
cells synapse
 Amacrine cells—where bipolar and ganglion cells
synapse
 Alter sensitivity of
retina; major role in
adjusting to dim or
bright light

Photoreceptor distribution
 Cones: ~6 million per eye
• Most dense at fovea centralis of macula—no rods
there
– Cone density directly correlates with visual acuity
(sharpness)
 Rods: ~125 million
per eye
• Maximum density
at periphery; few
cones there

A. What is the eye’s blind spot?

B. Compare rods with cones.
C. If you enter a dimly lit room, will you be able to
see clearly? Why or why not?
Learning Outcome: Describe the structure and

function of the retina’s layers of cells, and explain
the distribution of rods and cones and their relation
to visual acuity.

Module 15.11: Photoreception occurs in the
outer segment of rod and cone cells
Photoreceptors (rods and cones) detect photons

(basic units of light)
 Light energy—type of radiant energy that travels in
waves
 Our visible spectrum: 400–700 nm
Visual pigments transduce light
 Derived from rhodopsin (visual
purple); pigment in rods
 Have opsin (protein; determines
wavelength absorbed) and
retinal (pigment synthesized
from vitamin A)
Module 15.11: Photoreception occurs in the
outer segment of rod and cone cells
Visual pigments transduce light

 Derived from rhodopsin (visual purple); pigment in
rods
 Have opsin (protein; determines wavelength
absorbed) and retinal
(pigment synthesized
from vitamin A)

Module 15.11: Photoreception
Photoreceptor structure
 Pigmented epithelium
adjacent to
photoreceptors
• Absorbs photons not
absorbed by visual
pigments
• Phagocytizes old discs
shed from tip of outer
segment

(continued)
 Outer segment
• Flattened, membranous
discs containing visual
pigment
– Cones: plasma
membrane infoldings;
outer segment
tapers to blunt point
– Rods: discs are
separate structures;
outer segment forms
elongated cylinder
(continued)
 Inner segment
• Contains major
organelles—responsible
for all cell functions
other than
photoreception
• Each photoreceptor
synapses with a bipolar
cell

Color vision
 Rods all contain same opsin; responds to blue-
green wavelengths
 Three types of cones
1. Blue cones (16% of cones)
2. Green cones (10%)
3. Red cones (74%)

 Three cone types have different opsins; sensitive

to different wavelength ranges, with overlap
 If all three types are stimulated equally, we see
white

Color blindness—nonfunctional cones

 Inability to distinguish certain colors
 One or more types of cones are nonfunctional—
absent or do not make necessary visual pigment
 Most common type is red–green colorblindness—no
red cones; cannot distinguish between red and
green
 Often inherited
• 10 percent of males
• 0.67 percent of females
• 1 in 300,000—no pigment

A. Describe the structure of a photoreceptor.

B. How could a diet deficient in vitamin A affect
vision?
C. Identify the three types of cones.
Learning Outcome: Describe the structure of the

photoreceptors and how we are able to distinguish
colors.

Module 15.12: Photoreception involves activation,
bleaching, and reassembly of visual pigments
1. Resting state (in the dark)

 Chemically gated sodium ion channels of outer
segment stay open if cGMP present
 Inner segment continuously pumps sodium ions
out of cytosol

Module 15.12: Photoreception process
1. Resting state (in the dark) (continued)

 This movement of ions = dark current
 Keeps resting membrane potential about –40 mV
 Photoreceptor continually releases neurotransmitters
to bipolar cells

2. Retinal molecule in rhodopsin changes shape

(activation) from a bent 11-cis form to more
linear 11-trans form

3. Opsin activates transducin, a G protein bound

to disc membrane
 Transducin activates phosphodiesterase (PDE)

4. Phosphodiesterase breaks down cGMP,

inactivating gated sodium channels
 Sodium entry decreases

Active state
 Decreased sodium entry reduces dark current
 Membrane potential drops to –70 mV
(hyperpolarized)

Active state (continued)

 Hyperpolarization decreases neurotransmitter
release
 Decreased neurotransmitter signals bipolar cell that
the photoreceptor has absorbed a photon

The process of photoreception

Rhodopsin cannot respond to another photon until

original shape of retinal is regained
 Three step process
1. Bleaching
– Entire rhodopsin molecule first broken into retinal and
opsin
2. Retinal converted back to cis shape—requires ATP
3. Opsin and retinal are
reassembled as
rhodopsin

Module 15.12 Review
A. What are the two configurations of retinal?

B. Visual pigments undergo which three changes
during photoreception?
C. When during photoreception is ATP required?
Learning Outcome: Explain photoreception and

how visual pigments are activated.

Module 15.13: The visual pathways distribute
visual information from each eye to both
cerebral hemispheres
Visual pathways
 Photoreceptors to
bipolar cells to ganglion
cells
 ~1 million axons from
ganglion cell converge
at optic disc; head
toward diencephalon as
the optic nerve (II)

Module 15.13: Visual pathways
Visual pathways (continued)

 Two optic nerves (one from
each eye) reach diencephalon
at the optic chiasm
 From optic chiasm,
continue along optic
tracts
• About half of fibers
go to lateral geniculate
nucleus on same side
of brain; other half go
to opposite side


 Optic radiation = bundle of
projection fibers linking each
lateral geniculate
body with visual cortex,
in occipital cortex on
same side
 Collaterals from fibers
synapsing in lateral
geniculate bodies go to
subconscious
processing centers in
diencephalon and brainstem

 Pupillary reflexes and
others are triggered by
collaterals going to
superior colliculi

Perception of visual image reflects integration of

information arriving at visual cortex
 Depth perception = ability to judge depth or
distance by interpreting 3-D relationships
• Perceived by comparing relative positions of objects
within images received by both eyes
– Visual images from left and right eyes overlap
– Each eye receives slightly different image due to:
o Foveae centrales are 5–7.5 cm (2–3 in.) apart
o The nose and eye socket block view of opposite side

The visual pathway

Module 15.13 Review
A. Define optic radiation.

B. Where are visual images perceived?
Learning Outcome: Explain how the visual

pathways distribute information to their destinations
in the brain.

Module 15.14: Refractive problems result from
abnormalities in the cornea or lens or in the
shape of the eye
Emmetropia = normal vision
 When ciliary muscle is relaxed and lens flattened,
distant image is focused on retinal surface

Module 15.14: Refractive problems
Myopia = nearsightedness
 Focal distance too short; image
focuses in front of retina
• Eyeball too deep
• Resting curvature of lens too
great
 Correct with diverging (concave)
lens in front of eye
• Spreads light rays apart to shift
focus forward onto retina

Hyperopia = farsightedness
 Focal distance too long; image
focuses beyond retina
• Eyeball too shallow
• Lens too flat
 Correct with converging
(convex) lens in front of eye
• Provides additional refraction
to focus on retina

Surgical correction
 Photorefractive keratectomy (PRK)
 Computer-guided laser shapes cornea
• Removes 10–20 µm (< 10%) of cornea
• Completed in less than a minute
 Laser-assisted in-situ keratomileusis (LASIK)
 Interior corneal layers reshaped; covered by normal
corneal epithelium
 ~70% patients achieve normal vision
 ~10 million Americans have had it
 Immediate and long-term visual problems can occur

Module 15.14 Review
A. Define emmetropia.
B. Which type of lens would correct hyperopia?
Learning Outcome: Describe various refraction

problems associated with the cornea, lens, or
shape of the eye.

Section 3: Equilibrium and Hearing
Learning Outcomes
15.15 Describe the sensory receptors of the internal
ear.
15.16 Describe the structures of the external, middle,
and internal ear, and explain how they function.
15.17 Describe the structures and functions of the
bony labyrinth and membranous labyrinth.
15.18 Describe the functions of hair cells in the
semicircular ducts, utricle, and saccule.
15.19 Describe the structures and functions of the
spiral organ.

Section 3: Equilibrium and Hearing

15.20 Explain the anatomical and physiological basis
for pitch and volume sensations for hearing.
15.21 Trace the pathways for the sensations of
equilibrium and hearing to their respective
destinations in the brain.
15.22 Clinical Module: Describe age-related
disorders of olfaction, gustation, vision,
equilibrium, and hearing.

Module 15.15: Equilibrium and hearing involve
the internal ear
Comparison of receptors
 Olfactory receptors—specialized sensory neurons
 Gustatory receptors—communicate with sensory
neurons
 Photoreceptors—respond to light
 Both route information directly to the CNS
 All located in epithelia exposed to external
environment

Module 15.15: Internal ear sensory receptors
 Equilibrium and hearing receptors—isolated and

protected from external environment
 Located in internal ear
 Information integrated and organized locally;
forwarded to CNS

Hair cells = sensory receptors in internal ear

 Free surfaces covered with specialized nonmotile
processes
• Stereocilia—resemble long microvilli; 80–100 per
hair cell
• Kinocilium = single
large cilium

Hair cells are mechanoreceptors—sensitive to

contact/movement
 External force pushing on hair cell processes distorts
plasma membrane; alters neurotransmitter release
• Provides information about direction/strength of
stimulus
• Monitored by dendrites of sensory neurons

Complex 3-D structure in internal ear determines

what stimuli can reach hair cells in each region
 Hair cells in one region respond only to gravity or
acceleration
 Hair cells in other regions respond only to rotation or
to sound

Module 15.15 Review
A. Contrast the olfactory and gustatory sensory

receptors with those of equilibrium and hearing.
B. Describe the sensory receptors of the internal
ear.
C. What kind of stimuli can the internal ear sense?
Learning Outcome: Describe the sensory receptors

of the internal ear.

Module 15.16: The ear is divided into the
external ear, the middle ear, and the internal ear
External ear—collects/directs sound waves toward

middle ear
 Auricle—elastic cartilage
 External acoustic meatus—passageway in
temporal bone
• Ceruminous glands—
secrete waxy cerumen
(earwax); keeps foreign
objects out; slows growth
of microorganisms
• Hairs—trap debris

Module 15.16: Anatomy of the ear
Middle ear (tympanic cavity) = air-filled chamber

from tympanic membrane to auditory ossicles;
connects to pharynx by auditory tube
 Tympanic membrane (tympanum, eardrum) = thin,
semitransparent sheet that separates external ear
and middle ear
 Auditory ossicles =
three tiny bones; connect
tympanic membrane and
inner ear

Internal ear
 Contains sensory organs for hearing and equilibrium
 Receives amplified sound waves from middle ear
 Superficial contours established by layer of dense
bone = bony labyrinth

The anatomy of the ear

Middle ear
 Auditory tube
(pharyngotympanic
tube, eustachian tube)
• Connects middle ear
to nasopharynx
• Allows pressure
equalization across
tympanic membrane
• Can allow microorganisms into middle ear, causing
infection (otitis media)—can impair hearing, may
invade internal ear

Auditory ossicles
 Malleus (malleus, hammer)—attaches to tympanic
membrane
 Incus (incus, anvil)—attaches malleus to stapes
 Stapes (stapes, stirrup)—attached to oval window

Middle ear muscles

 Tensor tympani muscle connects malleus to
temporal bone
• Contraction stiffens tympanic membrane, reduces
vibration
• Innervated by mandibular
branch of trigeminal
nerve (V)

Middle ear muscles (continued)

 Stapedius muscle
• Connects stapes to posterior wall of middle ear
• Reduces stapes movement at oval window

Structures of the middle ear

Amplification and protection

 Sound waves vibrate tympanic membrane; convert
sound into mechanical movement
 Auditory ossicles conduct vibrations to internal ear
• Focuses sound on oval window and amplifies it
• Contractions of tensor tympani and stapedius
muscles protect tympanic membrane and ossicles
from violent movement under very noisy conditions

Module 15.6 Review
A. Name the three tiny bones located in the middle

ear, from lateral to medial.
B. What is the function of the auditory tube?
Learning Outcome: Describe the structures of the

external, middle, and internal ear, and explain how
they function.

Module 15.17: In the internal ear, the bony
labyrinth protects the membranous labyrinth
and its receptors
Bony labyrinth = shell of dense bone
surrounding/protecting membranous labyrinth
 Filled with perilymph = liquid similar to CSF;
between bony labyrinth and membranous labyrinth
 Three parts
1. Semicircular canals
2. Vestibule
3. Cochlea

Module 15.17: Labyrinths of the internal ear
Membranous labyrinth = collection of fluid-filled

tubes/chambers
 Houses receptors for hearing and equilibrium
 Contains fluid called endolymph

Three parts (semicircular canals, utricle, and

saccule are part of the vestibular complex, which
maintains equilibrium)
1. Semicircular ducts (within semicircular canals)
• Receptors stimulated by rotation of head
2. Within the vestibule—utricle and saccule
• Provide sensations of gravity and linear acceleration
3. Cochlear duct (within cochlea)
• Sandwiched between pair of perilymph-filled
chambers
• Resembles snail shell
• Receptors stimulated by sound
The anatomy of the internal ear

The membranous
labyrinth has four
parts. The
semicircular ducts,
utricle, and saccule
are part of the
vestibular complex,
which monitors
equilibrium. The
cochlear duct is
responsible for
hearing.

Module 15.7 Review
A. Identify the structures of the bony labyrinth.

B. How do the semicircular canals and the
semicircular ducts differ?
C. Describe the regional differences among the
receptor complexes in the membranous
labyrinth.
Learning Outcome: Describe the structures and

functions of the bony labyrinth and membranous
labyrinth.

Module 15.18: Hair cells in the semicircular
ducts respond to rotation; hair cells in the
utricle and saccule respond to gravity
and linear acceleration
Semicircular ducts
 Three ducts (anterior, posterior, lateral)—
continuous with utricle and filled with endolymph
 Ampulla = enlarged part of duct that houses
receptors

Module 15.18: Receptors for equilibrium
Semicircular ducts (continued)

 Ampullary crest = region in wall of ampulla with
receptors
 Ampullary cupula = gelatinous structure extending
through ampulla with kinocilia and stereocilia of hair
cells embedded in it

Head rotating in plane of a duct moves endolymph;

pushes ampullary cupula to side, distorting
receptor processes
 Movement in one direction causes stimulation;
opposite direction causes inhibition
 Ampullary cupula rebounds to normal position when
endolymph stops moving

Even complex angular movements can be

analyzed by movement of the three rotational
planes
 Horizontal rotation (“no”) stimulates lateral duct
receptors
 Nodding (“yes”) stimulates anterior duct receptors
 Tilting head to side stimulates posterior duct
receptors

Utricle and saccule

 Provide equilibrium sensations, whether body is
stationary or moving
 Connected by slender passageway continuous with
endolymphatic duct that ends in endolymphatic sac
• Sac projects into subarachnoid space
 Endolymphatic duct continuously secretes
endolymph; returns to general circulation at
endolymphatic sac

Utricle and saccule (continued)

 Utricle and saccule contain hair cells clustered in
maculae
• Macula of utricle senses horizontal movement
• Macula of saccule senses vertical movement
• Hair cell processes embedded in gelatinous otolithic
membrane
– Surface has densely packed
calcium carbonate
crystals (otoliths,
or “ear stones”)

Utricle and saccule (continued)

 Change in head position causes distortion of hair
cell processes in the maculae, sending signals to
the brain
• Head in upright position—otoliths
sit on top of otolithic membrane
in utricle
• Head in tilted position or with
linear movement—gravity pulls
on otoliths, shifts them to side
• Movement distorts hair cell
processes; stimulates
macular receptors

Module 15.18 Review
A. Define otoliths.
B. Cite the functions of sensory receptors in the
saccule and utricle.
Learning Outcome: Describe the functions of hair

cells in the semicircular ducts, utricle, and saccule.

Module 15.19: The cochlear duct contains the
hair cells of the spiral organ that function in
hearing
Cochlear duct (scala media)
 Filled with endolymph
Between two chambers
with perilymph
 Scala vestibuli
(vestibular duct)
 Scala tympani
(tympanic duct)
• Encased by bony labyrinth except at oval/round windows
 Interconnect at tip of cochlear, forming single long
chamber from oval window to round window
Module 15.19: Receptors for hearing
 Vestibular membrane separates cochlear

duct/scala vestibule
 Basilar membrane separates
cochlear duct from scala
tympani
 Hair cells for hearing located
in cochlear duct in the
spiral organ (organ
of Corti) on
basilar
membrane

Cross-sections of the cochlea

Cross-sectional anatomy of cochlea

 Scala vestibuli and scala tympani filled with
perilymph
 Cochlear duct filled with endolymph and contains
spiral organ (with receptors for hearing)
 Spiral ganglion—cell bodies of sensory neurons
monitoring adjacent hair cells in spiral organ
 Axons from spiral ganglion in
cochlear nerve of
vestibulocochlear
nerve (VIII)

Spiral organ
 Hair cells lack kinocilia
 Stereocilia are in contact with overlying tectorial
membrane
 Bulk of hair cell embedded in basilar membrane

Spiral organ (continued)

 Sound waves create pressure waves in perilymph
 Pressure waves cause basilar membrane to vibrate
up and down
 Vibrations of basilar membrane press stereocilia into
tectorial membrane, distorting them

Spiral organ (continued)

 Distortion triggers nerve impulse
 Sensory neurons relay signal through spiral
ganglion to cochlear branch of vestibulocochlear
nerve (VIII)

External and cross-sectional views of the
cochlea

Anatomy of the spiral organ

A pressure wave in the perilymph causes
movement of the hair cells and basilar
membrane.

Module 15.19 Review
A. Where is the spiral organ located?

B. Name the fluids found within the scala vestibuli,
scala tympani, and cochlear duct.
C. When the basilar membrane moves, what
happens to the hair cells of the spiral organ?
Learning Outcome: Describe the structures and

functions of the spiral organ.

Module 15.20: Sound waves lead to movement
of the basilar membrane in the process of
hearing
Hearing = perception of sound; sound = waves of
pressure
 In air, pressure wave causes alternating areas of
compressed/separated molecules
 Wavelength of sound =
distance between
adjacent wave crests
(peaks) or adjacent
troughs

Module 15.20: Physiology of hearing
Sound waves travel at the same speed (speed of

sound = 1235 km/h
 If frequency increases, wavelength decreases

Frequency = number of waves (cycles) passing

fixed point in given time
 Measured as cycles per second (cps);
units = hertz (Hz)
 Wavelength and frequency inversely related
 Wavelength and frequency inversely related
 Pitch = our perception
of frequency
• High frequency
(short wavelength)
= high pitch

Intensity (loudness) = amount of energy in sound

waves
 Amplitude of soundwave reflects amount of energy
(intensity)
Greater energy = larger amplitude = louder sound
 Measured in decibels (dB)


Sound waves and vibration

 Energy of sound waves is physical pressure
 Sound waves strike flexible object (i.e., tympanic
membrane); object responds
 At particular frequency and amplitude, object will
vibrate at same frequency = resonance
• Tympanic membrane resonates with sound waves,
generating movement of stapes at oval window
• Basilar membrane regions resonate at different
frequencies

For hearing:
 Stapes pushes on the oval window
• Inward movement causes distortion of basilar
membrane toward the round window
• Opposite action when stapes moves outward

For hearing: (continued)

 Flexibility of basilar membrane varies along its
length
• Different sound frequencies affect different parts of
the membrane
– Location of
vibration
interpreted
as pitch
– Number of
stimulated hair
cells interpreted
as volume

The role of the basilar membrane in hearing









Module 15.20 Review
A. Define decibel.
B. Beginning at the external acoustic meatus, list,
in order, the structures involved in hearing.
C. How would sound perception be affected if the
round window could not bulge out as a result of
increased perilymph pressure?
Learning Outcome: Explain the anatomical and

physiological basis for pitch and volume sensations
for hearing.

Module 15.21: The vestibulocochlear nerve
carries equilibrium and hearing sensations to
the brainstem
Equilibrium (balance)

Neural pathways for the sense of equilibrium





Module 15.21: Vestibulocochlear nerve function

Hearing
 Nerve signals for hearing are carried on the
cochlear nerve, which is part of the vestibulocohlear
nerve

Hearing
 Nerve signals for hearing are carried on the
cochlear nerve, which is part of the vestibulocohlear
nerve

Hearing (continued)
 Neural pathways for the sense of hearing

Hearing (continued)

Hearing (continued)

Hearing (continued)

Hearing (continued)
 Most auditory information from one cochlea is
projected to the auditory cortex on opposite side
 Some information from cochlea reaches auditory
cortex on its same side
• Aids in localizing sounds (left/right)
• Reduces functional impact of damage to one cochlea
or ascending pathway

Module 15.21 Review
A. Where are the hair cell receptors for equilibrium

located?
B. Which cranial nerves are involved with eye,
head, and neck movements?
C. What is your reflexive response to hearing a
loud noise, such as a firecracker?
Learning Outcome: Trace the pathways for the

sensations of equilibrium and hearing to their
respective destinations in the brain.

Module 15.22: Aging is associated with many
disorders of the special senses; trauma,
infection, and abnormal stimuli may cause
problems at any age
Olfaction disorders
 Head injury—damage to olfactory nerve (I)
 Age-related changes
• Olfactory receptors are regularly
replaced by stem cells, but
number declines with age
• Remaining receptors become
less sensitive

Module 15.22: Disorders of the special senses
Gustation disorders
 Problems with olfactory receptors—decreased smell
dulls taste
 Damaged taste buds—mouth infection, inflammation
 Damaged cranial nerves (VII, IX, X)—trauma or
compression
 Natural age-related changes

Vision disorders
 Senile cataract—lens loses transparency
• Natural consequence of aging; can be surgically
corrected
• Progresses—person needs more light to read; acuity
may decline to blindness
 Presbyopia—age-related farsightedness due to
loss of lens elasticity
(less accommodation
possible for close vision)

Equilibrium disorders
 Vertigo—false perception of spinning
• From conditions altering function of:
– Internal ear receptor complex
– Vestibular nerve (of vestibulocochlear nerve VIII)
– Sensory nuclei and CNS pathways
• Can be duo to vision problems or drug use (including
alcohol)

Vertigo (continued)
 Stimulated by anything that sets endolymph in
motion
 Motion sickness is most common cause
• Symptoms—headache, sweating, flushing of face,
nausea, vomiting

Hearing disorders
 Partial hearing deficit affects ~37.5 million in United
States
 Two types: conductive and sensorineural
Conductive hearing loss—problem conducting
sound waves
 Causes include
impacted earwax,
infection, perforated
tympanic membrane

Sensorineural hearing loss—damage to cochlea

or nerve pathways from internal ear to brain
 Causes include exposure to loud noise, head
trauma, and aging
 Age changes
• Tympanic membrane loses flexibility
• Articulations between auditory ossicles stiffen
• Round window may start to ossify

Module 15.22 Review
A. Which cranial nerves provide taste sensations

from the tongue?
B. What causes vertigo?
C. Identify two common classes of hearing-related
disorders.
Learning Outcome: Describe age-related disorders

of olfaction, gustation, vision, equilibrium, and
hearing.

Aand PCH 15 Lecture Pearson

Uploaded by

Copyright:

Available Formats

Aand PCH 15 Lecture Pearson

Uploaded by

Document Information

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Aand PCH 15 Lecture Pearson

Uploaded by

Copyright:

Available Formats

15

The Special Senses

© 2018 Pearson Education, Inc.

Copying slides from one slide set into another

1. Olfactory receptors are dendrites of specialized

© 2018 Pearson Education, Inc.

2. Receptors for taste, vision, equilibrium, hearing

© 2018 Pearson Education, Inc.

 Graded depolarization of the receptor cell triggers

© 2018 Pearson Education, Inc.

A. Where do the special senses originate?

Learning Outcome: Explain the roles of generator

© 2018 Pearson Education, Inc.

© 2018 Pearson Education, Inc.

© 2018 Pearson Education, Inc.

© 2018 Pearson Education, Inc.

© 2018 Pearson Education, Inc.

Odorants = dissolved chemicals that stimulate

© 2018 Pearson Education, Inc.

Olfactory reception process

© 2018 Pearson Education, Inc.

© 2018 Pearson Education, Inc.

© 2018 Pearson Education, Inc.

© 2018 Pearson Education, Inc.

Olfactory pathway to the cerebrum

© 2018 Pearson Education, Inc.

© 2018 Pearson Education, Inc.

© 2018 Pearson Education, Inc.

© 2018 Pearson Education, Inc.

© 2018 Pearson Education, Inc.

Learning Outcome: Describe the sensory organs of

© 2018 Pearson Education, Inc.

© 2018 Pearson Education, Inc.

Gustation = taste (continued)

© 2018 Pearson Education, Inc.

Gustation = taste (continued)

© 2018 Pearson Education, Inc.

Gustation = taste (continued)

© 2018 Pearson Education, Inc.

Gustation = taste (continued)

© 2018 Pearson Education, Inc.

© 2018 Pearson Education, Inc.

Four primary taste sensations: sweet, salty,

© 2018 Pearson Education, Inc.

Four primary taste sensations: sweet, salty,

© 2018 Pearson Education, Inc.

© 2018 Pearson Education, Inc.

Taste receptor sensitivity

© 2018 Pearson Education, Inc.

Learning Outcome: Describe the sensory organs of

© 2018 Pearson Education, Inc.

Gustatory reception stimulated by dissolved

© 2018 Pearson Education, Inc.

Two types of gustatory reception

2. Sweet, bitter, and umami receptors:

© 2018 Pearson Education, Inc.

Gustatory pathway—gustatory receptor cells to

© 2018 Pearson Education, Inc.

Gustatory pathway (continued)