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Chapter 15 Part A

The Special
Senses

PowerPoint® Lecture Slides


prepared by
Karen Dunbar Kareiva
© Annie Leibovitz/Contact Press Images
© 2016 Pearson Education, Inc. Ivy Tech Community College
Why This Matters

• Understanding the anatomy and physiology of


the eye helps to identify diseases such as
glaucoma

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Special Senses

• The sense of touch is one of the general


senses, mediated by general receptors (covered
in Chapter 13)
• Special senses of body include:
– Vision
– Taste
– Smell
– Hearing
– Equilibrium

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Special Senses (cont.)

• All use special sensory receptors, which are


distinct receptor cells localized in head region
– Not like modified nerves of general receptors

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Part 1 The Eye and Vision

• 70% of body’s sensory receptors are in eye


• Half of cerebral cortex is involved in visual
processing

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15.1 The Eye

• Small sphere; only one-sixth of surface visible


• Most of eye enclosed and protected by fat
cushion and bony orbit
• Consists of accessory structures and the
eyeball

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Figure 15.1a The eye and accessory structures.

Eyebrow

Eyelid

Eyelashes

Site where
conjunctiva
merges with
cornea

Palpebral
fissure

Lateral
commissure

Iris

Eyelid Pupil Sclera Lacrimal Medial


(covered by caruncle commissure
conjunctiva)
Surface anatomy of the right eye
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Accessory Structures of the Eye

• Accessory structures protect the eye and aid


eye function
• Structures include:
– Eyebrows
– Eyelids
– Conjunctiva
– Lacrimal apparatus
– Extrinsic eye muscles

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Accessory Structures of the Eye (cont.)

• Eyebrows
– Overlie supraorbital margins
– Function
• Shade eye from sunlight
• Prevent perspiration from reaching eye

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Accessory Structures of the Eye (cont.)

• Eyelids
– Also called palpebrae; thin, skin-covered folds
that protect eye anteriorly
– Separated at palpebral fissure (slit)
– Meet in corners at medial and lateral
commissures
– Lacrimal caruncle located at medial
commissure contains oil and sweat glands
– Tarsal plates: supporting connective tissue for
folds, as well as anchor orbicularis oculi and
levator palpebrae superioris muscles
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Accessory Structures of the Eye (cont.)

• Eyelids (cont.)
– Eyelids blink reflexively every 3–7 seconds
• Offers protection from foreign objects and spreads
secretions to moisten eye
– Eyelashes have follicles that are innervated
• Nerve endings initiate reflex blinking
– Lubricating glands associated with eyelids
• Tarsal (Meibomian) glands
– Modified sebaceous glands produce oily secretion that
lubricates lid and eye
• Ciliary glands between hair follicles are
– Modified sweat glands
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Figure 15.1b The eye and accessory structures.
Levator palpebrae
superioris muscle

Orbicularis
oculi muscle
Eyebrow
Tarsal plate
Palpebral
conjunctiva
Tarsal glands
Cornea

Palpebral
fissure

Eyelashes

Bulbar
conjunctiva

Conjunctival
sac

Orbicularis
oculi muscle

Lateral view; some structures shown in sagittal section


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Accessory Structures of the Eye (cont.)

• Conjunctiva
– Transparent mucous membrane that produces a
lubricating mucous secretion
– Palpebral conjunctiva: membrane that lines
underside of eyelids
– Bulbar conjunctiva: membrane that covers
white of eyes (not cornea)
• Small blood vessels found in this membrane; seen
easily in “bloodshot” eyes
– Conjunctival sac: space between palpebral and
bulbar conjunctiva
• Area where contact lens rests
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Accessory Structures of the Eye (cont.)

• Lacrimal apparatus
– Consists of lacrimal gland and ducts that drain
into nasal cavity
– Lacrimal gland is located in orbit above lateral
end of eye and secretes lacrimal secretion
(tears), a dilute saline solution containing mucus,
antibodies, and antibacterial lysozyme
– Blinking spreads tears toward medial
commissure, where they enter paired lacrimal
canaliculi via lacrimal puncta

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Accessory Structures of the Eye (cont.)

• Lacrimal apparatus (cont.)


– Tears then drain into lacrimal sac and
nasolacrimal duct, which empties into nasal
cavity

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Figure 15.2 The lacrimal apparatus.

Lacrimal sac
Lacrimal gland
Excretory ducts
of lacrimal gland

Lacrimal punctum

Lacrimal canaliculus

Nasolacrimal duct

Inferior meatus
of nasal cavity

Nostril

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Accessory Structures of the Eye (cont.)

• Extrinsic eye muscles


– Six straplike extrinsic eye muscles
• Originate from bony orbit and insert on eyeball
• Enable eye to follow moving objects, maintain shape
of eyeball, and hold it in orbit
– Four rectus muscles originate from common
tendinous ring; names indicate movements
• Superior, inferior, lateral, and medial rectus
– Two oblique muscles move eye in vertical plane
and rotate eyeball
• Superior and inferior oblique muscles

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Figure 15.3a Extrinsic eye muscles.

Superior oblique
tendon

Superior oblique
muscle

Superior rectus
muscle

Lateral rectus
muscle

Inferior rectus Inferior oblique


muscle muscle

Lateral view of the right eye


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Figure 15.3b Extrinsic eye muscles.

Trochlea

Superior oblique
tendon
Axis of
Superior oblique rotation
muscle of eye

Superior rectus Inferior


muscle rectus muscle

Medial
rectus muscle

Lateral
rectus muscle

Common
tendinous ring

Superior view of the right eye


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Figure 15.3c Extrinsic eye muscles.

Trochlea
Superior
rectus Superior
oblique
Lateral Medial
rectus rectus

Inferior Inferior
oblique rectus

Anterior view of the right eye

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Figure 15.3d Extrinsic eye muscles.

Muscle Action Controlling cranial nerve

Lateral rectus Moves eye laterally VI (abducens)


Medial rectus Moves eye medially III (oculomotor)
Superior rectus Elevates eye and turns it medially III (oculomotor)
Inferior rectus Depresses eye and turns it medially III (oculomotor)
Inferior oblique Elevates eye and turns it laterally III (oculomotor)
Superior oblique Depresses eye and turns it laterally IV (trochlear)

Summary of muscle actions and innervating cranial nerves

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Clinical – Homeostatic Imbalance 15.1

• An infected tarsal gland results in an unsightly


cyst called a chalazion
• Inflammation of any of the smaller sebaceous
glands is called a sty

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Clinical – Homeostatic Imbalance 15.2

• Conjunctivitis: inflammation of the conjunctiva


resulting in reddened, irritated eyes
• Pinkeye: conjunctival infection caused by
bacteria or viruses
– Highly contagious

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Clinical – Homeostatic Imbalance 15.3

• Nasal cavity mucosa is continuous with mucosa


of lacrimal duct system, so a cold or nasal
inflammation often causes lacrimal mucosa to
swell
• Swelling constricts the ducts and prevents tears
from draining, causing “watery” eyes

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Clinical – Homeostatic Imbalance 15.4

• Diplopia (double vision): occurs when


movements of external muscles of two eyes are
not perfectly coordinated
– Person cannot properly focus images of same
area of the visual field from each eye, so sees
two images instead of one
– Can result from paralysis, extrinsic muscle
weakness, or neurological disorders

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Clinical – Homeostatic Imbalance 15.4

• Strabismus (“cross-eye”): congenital


weakness of external eye muscles
– Eye rotates medially or laterally
– Eyes may alternate focusing on objects, or only
controllable eye is used
• Brain begins to disregard inputs from deviant eye,
which can become functionally blind if not treated
early

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Structure of the Eyeball

• Wall of eyeball contains three layers


– Fibrous layer
– Vascular layer
– Inner layer
• Internal cavity filled with fluids called humors
• Lens separates internal cavity into anterior and
posterior segments

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Figure 15.4a Internal structure of the eye (sagittal section).

Ora serrata

Ciliary body Sclera


Ciliary zonule Choroid
(suspensory
ligament) Retina
Cornea
Macula lutea
Iris
Fovea centralis
Pupil
Optic nerve

Anterior
segment
(contains
aqueous humor)
Lens
Scleral venous sinus Central artery and
vein of the retina
Posterior segment Optic disc
(contains vitreous humor) (blind spot)

Diagrammatic view. The vitreous humor is illustrated only in the bottom part of the eyeball.

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Structure of the Eyeball (cont.)

• Fibrous layer
– Outermost layer; dense avascular connective
tissue
– Two regions: sclera and cornea
1. Sclera
– Opaque posterior region
– Protects and shapes eyeball
– Anchors extrinsic eye muscles
– Posteriorly, where optic nerve exits, sclera is
continuous with dura mater of brain

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Structure of the Eyeball (cont.)

2. Cornea
– Transparent anterior one-sixth of fibrous layer
» Forms clear window that lets light enter and bends
light as it enters eye
– Epithelium covers both surfaces
» Outer surface protects from abrasions
» Inner layer, corneal endothelium, contains sodium
pumps that help maintain clarity of cornea
– Numerous pain receptors contribute to blinking and
tearing reflexes

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Structure of the Eyeball (cont.)

• Vascular layer
– Middle pigmented layer of eye, also called uvea
– Three regions: choroid, ciliary body, and iris
1. Choroid region
– Posterior portion of uvea
– Supplies blood to all layers of eyeball
– Brown pigment absorbs light to prevent scattering of
light, which would cause visual confusion

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Structure of the Eyeball (cont.)

2. Ciliary body
– Anteriorly, choroid becomes ciliary body
– Thickened ring of tissue surrounding lens
– Consists of smooth muscle bundles, ciliary muscles,
that control shape of lens
– Capillaries of ciliary processes secrete fluid for
anterior segment of eyeball
– Ciliary zonule (suspensory ligament) extends from
ciliary processes to lens
» Holds lens in position

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Structure of the Eyeball (cont.)

3. Iris
– Colored part of eye that lies between cornea and lens,
continuous with ciliary body
– Pupil: central opening that regulates amount of light
entering eye
» Close vision and bright light cause sphincter
pupillae (circular muscles) to contract and pupils to
constrict; parasympathetic control
» Distant vision and dim light cause dilator pupillae
(radial muscles) to contract and pupils to dilate;
sympathetic control
» Changes in emotional state—pupils dilate when
subject matter is appealing or requires problem-
solving skills
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Figure 15.4a Internal structure of the eye (sagittal section).

Ora serrata

Ciliary body Sclera


Ciliary zonule Choroid
(suspensory
ligament) Retina
Cornea
Macula lutea
Iris
Fovea centralis
Pupil
Optic nerve

Anterior
segment
(contains
aqueous humor)
Lens
Scleral venous sinus Central artery and
vein of the retina
Posterior segment Optic disc
(contains vitreous humor) (blind spot)

Diagrammatic view. The vitreous humor is illustrated only in the bottom part of the eyeball.

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Figure 15.4b Internal structure of the eye (sagittal section).

Ciliary body Vitreous humor


Ciliary in posterior
processes segment

Iris Retina
Margin Choroid
of pupil
Anterior Sclera
segment
Fovea centralis
Lens
Cornea Optic nerve

Ciliary zonule Optic disc


(suspensory
ligament)
Photograph of the human eye.

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Figure 15.5 Pupil constriction and dilation.

Parasympathetic + Sympathetic +

Sphincter pupillae Iris (two muscles) Dilator pupillae


muscle contracts: • Sphincter pupillae muscle contracts:
Pupil constricts • Dilator pupillae Pupil dilates
(size decreases). (size increases).

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Structure of the Eyeball (cont.)

• Inner layer (retina)


– Retina originates as an outpocketing of brain
– Contains:
• Millions of photoreceptor cells that transduce light
energy
• Neurons
• Glial cells
– Delicate two-layered membrane
• Outer pigmented layer
• Inner neural layer

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Figure 15.6a Microscopic anatomy of the retina.

Neural layer of retina

Pigmented
Pathway layer of
of light retina
Choroid

Optic disc Sclera

Central artery
and vein of retina
Optic
nerve

Posterior aspect of the eyeball

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Structure of the Eyeball (cont.)

• Inner layer (retina) (cont.)


– Pigmented layer of the retina
• Single-cell-thick lining next to choroid
• Extends anteriorly, covering ciliary body and iris
• Functions:
– Absorbs light and prevents its scattering
– Phagocytizes photoreceptor cell fragments
– Stores vitamin A

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Structure of the Eyeball (cont.)

• Inner layer (retina) (cont.)


– Neural layer of the retina
• Transparent layer that runs anteriorly to margin of
ciliary body
– Anterior end has serrated edges called ora serrata
• Composed of three main types of neurons
– Photoreceptors, bipolar cells, ganglion cells
• Signals spread from photoreceptors to bipolar cells to
ganglion cells
• Ganglion cell axons exit eye as optic nerve

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Structure of the Eyeball (cont.)

• Inner layer (retina) (cont.)


– Neural layer of the retina (cont.)
• Optic disc
– Site where optic nerve leaves eye
– Lacks photoreceptors, so referred to as blind spot
• Retina has quarter-billion photoreceptors that are one
of two types:
– Rods
– Cones

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Figure 15.6a Microscopic anatomy of the retina.

Neural layer of retina

Pigmented
Pathway layer of
of light retina
Choroid

Optic disc Sclera

Central artery
and vein of retina
Optic
nerve

Posterior aspect of the eyeball

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Structure of the Eyeball (cont.)

• Inner layer (retina) (cont.)


– Rods
• Dim light, peripheral vision receptors
• More numerous and more sensitive to light than cones
• No color vision or sharp images
• Numbers greatest at periphery

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Structure of the Eyeball (cont.)

• Inner layer (retina) (cont.)


– Cones
• Vision receptors for bright light
• High-resolution color vision
• Macula lutea area at posterior pole lateral to blind
spot
– Contains mostly cones
• Fovea centralis: tiny pit in center of macula lutea that
contains all cones, so is region with best visual acuity
– Eye movement allows us to focus in on object so that
fovea can pick it up

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Figure 15.6b Microscopic anatomy of the retina.

Ganglion
cells Photoreceptors
Bipolar • Rod
Axons cells • Cone
of
ganglion
cells

Horizontal
cell

Amacrine cell
Pathway of signal output
Pigmented
Pathway of light layer of retina
Cells of the neural layer of the retina
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Figure 15.6c Microscopic anatomy of the retina.

Choroid
Outer segments
of rods and cones
Nuclei of
ganglion
cells

Axons of Nuclei
ganglion cells of bipolar
Nuclei of Pigmented
cells
rods and layer of retina
cones
Photomicrograph of retina
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Structure of the Eyeball (cont.)

• Inner layer (retina) (cont.)


– Two sources of blood supply to retina
• Choroid supplies outer third (photoreceptors)
• Central artery and vein of retina supply inner
two-thirds
– Enter/exit eye in center of optic nerve
– Vessels are visible in living person

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Figure 15.7 Part of the posterior wall (fundus) of the right eye as seen with an ophthalmoscope.

Central
artery
and vein
emerging
from the
optic disc

Optic disc

Macula
lutea

Retina
Lateral Medial

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Clinical – Homeostatic Imbalance 15.5

• Retinal detachment: condition where pigmented


and neural layers separate (detach), allowing
jellylike vitreous humor to seep between them
• Can lead to permanent blindness
• Usually happens when retina is torn during
traumatic blow to head or sudden stopping of
head during movement (example: bungee
jumping)

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Clinical – Homeostatic Imbalance 15.5

• Symptom described by victims as “curtain being


drawn across the eye” and/or sootlike spots or
light flashes
• Treatment: reattachment of retina with laser
surgery

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Structure of the Eyeball (cont.)

• Internal chambers and fluids


– The lens and ciliary zonule separate eye into two
segments
1. Posterior segment
2. Anterior segment

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Structure of the Eyeball (cont.)

• Internal chambers and fluids (cont.)


– Posterior segment
• Contains vitreous humor, a fluid that:
– Transmits light
– Supports posterior surface of lens
– Holds neural layer of retina firmly against pigmented
layer
– Contributes to intraocular pressure
• Vitreous humor forms in embryo and lasts whole
lifetime

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Structure of the Eyeball (cont.)

• Internal chambers and fluids (cont.)


– Anterior segment
• Iris divides anterior segment into two chambers:
– Anterior chamber—between cornea and iris
– Posterior chamber—between iris and lens
• Entire segment contains aqueous humor, a plasma
like fluid continuously formed (unlike vitreous humor)
by capillaries of ciliary processes
– Drains via scleral venous sinus (canal of Schlemm) at
sclera-cornea junction
– Supplies nutrients and oxygen mainly to lens and
cornea but also to retina, and removes wastes

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Figure 15.8 Circulation of aqueous humor. Slide 2

Iris
Posterior
Lens epithelium
segment
Cornea Lens
Lens (contains
vitreous
Cornea humor)
Corneal epithelium
Corneal endothelium
Aqueous humor

Anterior chamber
Anterior Ciliary zonule
segment Posterior chamber
(suspensory
(contains ligament)
aqueous
humor) 1
Scleral venous Ciliary
sinus
1 Aqueous humor forms by processes
filtration from the capillaries in Corneoscleral Ciliary
the ciliary processes. junction body
Ciliary
muscle
Bulbar
conjunctiva

Sclera

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Figure 15.8 Circulation of aqueous humor. Slide 3

Iris
Posterior
Lens epithelium
segment
Cornea Lens
Lens (contains
vitreous
Cornea humor)
2
Corneal epithelium
Corneal endothelium
Aqueous humor

Anterior chamber
Anterior Ciliary zonule
segment Posterior chamber
(suspensory
(contains ligament)
aqueous
humor) 1
Scleral venous Ciliary
sinus
1 Aqueous humor forms by processes
filtration from the capillaries in Corneoscleral Ciliary
the ciliary processes. junction body
Ciliary
muscle
Bulbar
2 Aqueous humor flows from
conjunctiva
the posterior chamber through the
pupil into the anterior chamber. Sclera
Some also flows through the
vitreous humor (not shown).

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Figure 15.8 Circulation of aqueous humor. Slide 4

Iris
Posterior
Lens epithelium
segment
Cornea Lens
Lens (contains
vitreous
Cornea humor)
2
Corneal epithelium
Corneal endothelium
Aqueous humor

Anterior chamber
Anterior Ciliary zonule
segment Posterior chamber
(suspensory
(contains ligament)
aqueous
humor) 3 1
Scleral venous Ciliary
sinus
1 Aqueous humor forms by processes
filtration from the capillaries in Corneoscleral Ciliary
the ciliary processes. junction body
Ciliary
muscle
Bulbar
2 Aqueous humor flows from
conjunctiva
the posterior chamber through the
pupil into the anterior chamber. Sclera
Some also flows through the
vitreous humor (not shown).

3 Aqueous humor is reabsorbed


into the venous blood by the scleral
venous sinus.

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Clinical – Homeostatic Imbalance 15.6

• Glaucoma: condition in which drainage of


aqueous humor is blocked, causing fluid to back
up and increase pressure within eye
• Pressures may increase to dangerous levels
and compress retina and optic nerve, leading to
blindness
• Symptoms: few early signs, but late signs
include seeing halos around lights and blurred
vision

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Clinical – Homeostatic Imbalance 15.6

• Detection: intraocular pressure determined by


directing puff of air at cornea and measuring
amount of corneal deformation
– Test should be done yearly after age 40
• Treatment: eye drops that increase rate of
aqueous humor drainage or decrease its
production; laser therapy or surgery

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Structure of the Eyeball (cont.)

• Lens
– Biconvex, transparent, flexible, and avascular
– Changes shape to precisely focus light on retina
– Two regions:
1. Lens epithelium: anterior region of cuboidal cells
that differentiate into lens fiber cells
2. Lens fibers: form bulk of lens and are filled with
transparent protein crystallin
– Lens fibers are continually added, so lens
becomes more dense, convex, and less elastic
with age
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Clinical – Homeostatic Imbalance 15.7

• Clouding of lens
– Consequence of aging, diabetes mellitus, heavy
smoking, frequent exposure to intense sunlight
– Some congenital
– Crystallin proteins clump
– Vitamin C increases cataract formation
– Lens can be replaced surgically with artificial
lens

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Figure 15.9 Photograph of a cataract.

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15.2 Focusing and Light

Overview: Light and Optics


• Wavelength and color
• Electromagnetic radiation: all energy waves, from long
radio waves to short X rays; visible light occupies a small
portion in the middle of the spectrum
• Light has wavelengths between 400 and 700 nm
• Eyes respond only to visible light

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Figure 15.10a The electromagnetic spectrum and photoreceptor sensitivities.

(10 9 nm=)
10−5 nm 10−3 nm 1 nm 103 nm 106 nm 1m 103 m

Gamma
X rays UV Infrared Micro- Radio waves
rays waves

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Overview: Light and Optics

• Wavelength and color (cont.)


• Light: packets of energy (photons or quanta) that
travel in wavelike fashion at high speeds
• When visible light passes through spectrum, it is
broken up into bands of colors (rainbow)
• Red wavelengths are longest and have lowest energy,
and violet are shortest and have most energy
• Color that eye perceives is a reflection of that
wavelength
• Grass is green because it absorbs all colors except green
• White reflects all colors, and black absorbs all colors

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Figure 15.10b The electromagnetic spectrum and photoreceptor sensitivities.
Visible light

Light absorption (percent of maximum)


Blue Green Red
cones Rods cones cones
(420 nm) (500 nm) (530 nm) (560 nm)

100

50

0
400 450 500 550 600 650 700
Wavelength (nm)

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Overview: Light and Optics (cont.)

• Refraction and lenses


• Refraction: bending of light rays
• Due to change in speed of light when it passes from one
transparent medium to another and path of light is at an
oblique angle
• Example: from liquid to air
• Lenses of eyes can also refract light because they are
curved on both sides
• Convex: thicker in center than at edges
• Concave: thicker at edges than in center

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Figure 15.11 Refraction.

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Overview: Light and Optics (cont.)

• Refraction and lenses (cont.)


• Convex lenses bend light passing through it, so that
rays converge at focal point
• Image formed at focal point is upside-down and reversed
from left to right
• Concave lenses disperse light, preventing light from
being focused

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Figure 15.12 Light is focused by a convex lens.

Point sources Focal points

Focusing of two points of light.

The image is inverted—upside down and reversed.


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Focusing Light on the Retina

• Pathway of light entering eye: cornea, aqueous


humor, lens, vitreous humor, entire neural layer of
retina, and finally photoreceptors
• Light is refracted three times along path:
(1) entering cornea, (2) entering lens, and
(3) leaving lens
• Majority of refractory power is in cornea; however, it
is constant and cannot change focus

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Focusing Light on the Retina (cont.)

• Lens is able to adjust its curvature to allow for fine


focusing
• Can focus for distant vision and for close vision

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Figure 15.13a Focusing for distant and close vision.

The ciliary muscle and the ciliary zonule focus an


image by changing the shape of the lens.
• They are arranged sphincterlike around the lens.
• Ciliary muscle contraction loosens the ciliary zonule fibers
and relaxation tightens them.

View

Ciliary muscle
Lens

Ciliary zonule
(suspensory ligament)

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Focusing Light on the Retina (cont.)

• Focusing for distant vision


• Eyes are best adapted for distant vision
• Far point of vision: distance beyond which no
change in lens shape is needed for focusing
• 20 feet for emmetropic (normal) eye
• Cornea and lens focus light precisely on retina at this
distance
• Ciliary muscles are completely relaxed in distance
vision, which causes a pull on ciliary zonule; as a
result, lenses are stretched flat

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Figure 15.13b Focusing for distant and close vision.

The lens flattens for distant vision.


Sympathetic input relaxes the ciliary muscle. This tightens
the ciliary zonule and flattens the lens.

Relaxed ciliary muscle


Tightened ciliary zonule
Flattened lens

Nearly parallel rays


from distant object Image

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Focusing Light on the Retina (cont.)

• Focusing for close vision


• Light from close objects (<6 m) diverges as
approaches eye
• Requires eye to make active adjustments using
three simultaneous processes:
1. Accommodation of the lenses
• Changing lens shape to increase refraction
• Near point of vision
• Closest point on which the eye can focus
• Presbyopia: loss of accommodation over age 50

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Focusing Light on the Retina (cont.)

2. Constriction of the pupils


• Accommodation pupillary reflex involves constriction of
pupils to prevent most divergent light rays from entering
eye
• Mediated by parasympathetic nervous system
3. Convergence of the eyeballs
• Medial rotation of eyeballs causes convergence of eyes
toward object being viewed
• Controlled by somatic motor neuron innervation on medial
rectus muscles

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Figure 15.13c Focusing for distant and close vision.

The lens bulges for close vision.


Parasympathetic input contracts the ciliary muscle. This
loosens the ciliary zonule and allows the lens to bulge.

Contracted ciliary muscle


Loosened ciliary zonule
Bulging lens

Divergent rays Image


from close object

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Clinical – Homeostatic Imbalance 15.8

• Problems associated with refraction related to


eyeball shape:
• Myopia (nearsightedness)
• Eyeball is too long, so focal point is in front of retina
• Corrected with a concave lens
• Hyperopia (farsightedness)
• Eyeball is too short, so focal point is behind retina
• Corrected with a convex lens
• Astigmatism
• Unequal curvatures in different parts of cornea or lens
• Corrected with cylindrically ground lenses or laser
procedures

© 2016 Pearson Education, Inc.


Figure 15.14-1 Problems of refraction.

Emmetropic eye (normal)


Focal point is on retina.
Focal
plane

Myopic eye (nearsighted: eyeball too long)


Uncorrected
Focal point is in front
of retina.
Eyeball
too long

Corrected
Concave lens moves focal
point further back.

© 2016 Pearson Education, Inc.


Figure 15.14-2 Problems of refraction.

Emmetropic eye (normal)


Focal point is on retina.
Focal
plane

Hyperopic eye (farsighted: eyeball too short)


Uncorrected
Focal point is behind
retina.
Eyeball
too short

Corrected
Convex lens moves focal
point forward.

© 2016 Pearson Education, Inc.


15.3 Phototransduction

Functional Anatomy of Photoreceptors


• Photoreceptors (rods and cones) are modified
neurons that resemble upside-down epithelial
cells
• Consists of cell body, synaptic terminal, and
two segments:
• Outer segment: light-receiving region
• Contains visual pigments (photopigments) that change
shape as they absorb light
• Inner segment of each joins cell body
• Inner segment is connected via cilium to outer segment
© 2016 Pearson Education, Inc.
Functional Anatomy of Photoreceptors (cont.)

• Cell body is connected to synaptic terminal via


inner fibers
• Plasma membrane of outer segment folds back to
form many discs
• Photopigments are embedded in discs

© 2016 Pearson Education, Inc.


Functional Anatomy of Photoreceptors (cont.)

• Photoreceptors are vulnerable to damage


• Degenerate if retina detached
• Destroyed by intense light
• Vision is maintained because outer segment is
renewed every 24 hours
• Tips fragment off and are phagocytized

© 2016 Pearson Education, Inc.


Figure 15.15a Photoreceptors of the retina.
Process
of bipolar
cell

Synaptic
Inner terminals
fibers
Rod cell
Rod body
cell
body
Nuclei
Cone
cell
body
Mitochondria
Outer
fiber Connecting
cilia

Inner segment
Apical

Outer segment
microvillus

Pigmented layer

Discs
containing
visual pigments

Discs being
phagocytized

Pigment
cell
Melanin
nucleus
granules

Basal lamina
(border with
choroid)

The outer segments of rods and cones


are embedded in the pigmented layer
of the retina.
© 2016 Pearson Education, Inc.
Comparing Rod and Cone Vision

• Rods are very sensitive to light, making them


best suited for night vision and peripheral
vision
• Contain a single pigment, so vision is perceived in
gray tones only
• Pathways converge, causing fuzzy, indistinct
images
• As many as 100 rods may converge into one ganglion

© 2016 Pearson Education, Inc.


Comparing Rod and Cone Vision (cont.)

• Cones have low sensitivity, so require bright


light for activation
• React more quickly than rods
• Have one of three pigments, which allow for vividly
colored sight
• Nonconverging pathways result in detailed,
high-resolution vision
• Some cones have their own ganglion cell, so brain can
put together accurate, high-acuity resolution images

© 2016 Pearson Education, Inc.


Table 15.1 Comparison of Rods and Cones

© 2016 Pearson Education, Inc.


Clinical – Homeostatic Imbalance 15.9

• Color blindness: lack of one or more cone


pigments
• Inherited as an X-linked condition, so more
common in males
• As many as 8–10% of males have some form
• The most common type is red-green, in which
either red cones or green cones are absent
• Depending on which cone is missing, red can
appear green, or vice versa
• Rely on different shades to get cues of color

© 2016 Pearson Education, Inc.


Visual Pigments

• Retinal: key light-absorbing molecule that


combines with one of four proteins (opsins) to
form visual pigments
• Synthesized from vitamin A
• Four opsins are rhodopsin (found in rods only), and
three found in cones: green, blue, red (depending
on wavelength of light they absorb)
• Cone wavelengths do overlap, so same wavelength
may trigger more than one cone, enabling us to see
variety of hues of colors
• Example: yellow light stimulates red and green cones, but
if more red are triggered, we see orange

© 2016 Pearson Education, Inc.


Visual Pigments (cont.)

• Retinal isomers are different 3-D forms


• Retinal is in a bent form in dark, but when pigment
absorbs light, it straightens out
• Bent form called 11-cis-retinal
• Straight form called all-trans-retinal
• Conversion of bent to straight initiates reactions that lead
to electrical impulses along optic nerve

© 2016 Pearson Education, Inc.


Figure 15.15b Photoreceptors of the retina.

Rod discs

Visual
pigment
consists of
• Retinal
• Opsin

Rhodopsin, the visual pigment in rods,


is embedded in the membrane that
forms discs in the outer segment.
© 2016 Pearson Education, Inc.
Phototransduction

• Phototransduction: process by which pigment


captures photon of light energy, which is
converted into a graded receptor potential
• Capturing light
• Deep purple pigment of rods is rhodopsin
• Arranged in rod’s outer segment
• Three steps of rhodopsin formation and breakdown:
• Pigment synthesis, pigment bleaching, and pigment
regeneration
• Similar process in cones, but different types of
opsins and cones require more intense light

© 2016 Pearson Education, Inc.


Phototransduction (cont.)

• Capturing light (cont.)


• Pigment synthesis
• Opsin and 11-cis retinal combine to form rhodopsin in dark
• Pigment bleaching
• When rhodopsin absorbs light, 11-cis isomer of retinal
changes to all-trans isomer
• Retinal and opsin separate (rhodopsin breakdown)
• Pigment regeneration
• All-trans retinal converted back to 11-cis isomer
• Rhodopsin is regenerated in outer segments

© 2016 Pearson Education, Inc.


Figure 15.16 The formation and breakdown of rhodopsin. Slide 2
11-cis-retinal

2H+

1 Pigment synthesis: Oxidation


11-cis-retinal, derived
from vitamin A, is Vitamin A
combined with opsin 11-cis-retinal
Rhodopsin
to form rhodopsin. Reduction

2H+

Dark Light

Opsin
and

All-trans-retinal

All-trans-retinal

© 2016 Pearson Education, Inc.


Figure 15.16 The formation and breakdown of rhodopsin. Slide 3
11-cis-retinal

2H+

1 Pigment synthesis: Oxidation


11-cis-retinal, derived
from vitamin A, is Vitamin A
combined with opsin 11-cis-retinal
Rhodopsin
to form rhodopsin. Reduction

2 Pigment bleaching:
2H+
Light absorption by
rhodopsin triggers a
rapid series of steps in
Dark Light which retinal changes
shape (11-cis to all-
trans) and eventually
releases from opsin.

Opsin
and

All-trans-retinal

All-trans-retinal

© 2016 Pearson Education, Inc.


Figure 15.16 The formation and breakdown of rhodopsin. Slide 4
11-cis-retinal

2H+

1 Pigment synthesis: Oxidation


11-cis-retinal, derived
from vitamin A, is Vitamin A
combined with opsin 11-cis-retinal
Rhodopsin
to form rhodopsin. Reduction

2 Pigment bleaching:
2H+
Light absorption by
rhodopsin triggers a
rapid series of steps in
Dark Light which retinal changes
3 Pigment regeneration: shape (11-cis to all-
Enzymes slowly convert trans) and eventually
all-trans-retinal to its releases from opsin.
11-cis form in cells of the
pigmented layer; requires
ATP.

Opsin
and

All-trans-retinal

All-trans-retinal

© 2016 Pearson Education, Inc.


Phototransduction (cont.)

• Light transduction reactions


• Light-activated rhodopsin activates G protein
transducin
• Transducin activates PDE, which breaks down
cyclic GMP (cGMP)
• In dark, cGMP holds cation channels of outer
segment open
• Na+ and Ca2+ enter and depolarize cell
• In light cGMP breaks down, channels close, cell
hyperpolarizes
• Hyperpolarization is signal for vision!

© 2016 Pearson Education, Inc.


Figure 15.17 Events of phototransduction. Slide 2

G protein signaling mechanisms


are like a molecular relay race.

Light (1st Receptor G protein Enzyme 2nd


messenger) messenger

1 Retinal absorbs light


and changes shape. Visual
pigment activates.
Ca2+ Ca2+
Visual Na+
pigment Na+
Light

cGMP cGMP

GMP
11-cis-retinal

Transducin
(a G protein)

© 2016 Pearson Education, Inc.


Figure 15.17 Events of phototransduction. Slide 3

G protein signaling mechanisms


are like a molecular relay race.

Light (1st Receptor G protein Enzyme 2nd


messenger) messenger

1 Retinal absorbs light


and changes shape. Visual
pigment activates.
Ca2+ Ca2+
Visual Na+
pigment Na+
All-trans-retinal
Light

cGMP cGMP

GMP
11-cis-retinal

Transducin 2 Visual pigment


(a G protein) activates transducin
(G protein).

© 2016 Pearson Education, Inc.


Figure 15.17 Events of phototransduction. Slide 4

G protein signaling mechanisms


are like a molecular relay race.

Light (1st Receptor G protein Enzyme 2nd


messenger) messenger

1 Retinal absorbs light


and changes shape. Visual
pigment activates.
Phosphodiesterase (PDE) Ca2+ Ca2+
Visual Na+
pigment Na+
All-trans-retinal
Light

cGMP cGMP

GMP
11-cis-retinal

Transducin 2 Visual pigment 3 Transducin


(a G protein) activates transducin activates
(G protein). phosphodiesterase
(PDE).

© 2016 Pearson Education, Inc.


Figure 15.17 Events of phototransduction. Slide 5

G protein signaling mechanisms


are like a molecular relay race.

Light (1st Receptor G protein Enzyme 2nd


messenger) messenger

1 Retinal absorbs light


and changes shape. Visual
pigment activates.
Phosphodiesterase (PDE) Ca2+ Ca2+
Visual Na+
pigment Na+
All-trans-retinal
Light

cGMP cGMP

GMP
11-cis-retinal

Transducin 2 Visual pigment 3 Transducin 4 PDE converts


(a G protein) activates transducin activates cGMP into GMP,
(G protein). phosphodiesterase causing cGMP
(PDE). levels to fall.

© 2016 Pearson Education, Inc.


Figure 15.17 Events of phototransduction. Slide 6

G protein signaling mechanisms


are like a molecular relay race.

Light (1st Receptor G protein Enzyme 2nd


messenger) messenger

1 Retinal absorbs light


and changes shape. Visual
pigment activates.
Phosphodiesterase (PDE) Ca2+ Ca2+
Visual Na+
pigment Na+
All-trans-retinal
Light

cGMP cGMP

cGMP-gated cGMP-gated
GMP cation channel cation channel
open in dark closed in light
11-cis-retinal

Transducin 2 Visual pigment 3 Transducin 4 PDE converts 5 As cGMP levels fall,


(a G protein) activates transducin activates cGMP into GMP, cGMP-gated cation
(G protein). phosphodiesterase causing cGMP channels close, resulting
(PDE). levels to fall. in hyperpolarization.

© 2016 Pearson Education, Inc.


Information Processing in the Retina

• Photoreceptors and bipolar cells generate only


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

© 2016 Pearson Education, Inc.


Figure 15.18-1 Signal transmission in the retina. Slide 2
Below, we look at a tiny column of retina.
The outer segment of the rod, closest to
the back of the eye and farthest from the
incoming light, is at the top.

In the dark

Light
1 cGMP-gated channels Na+
open, allowing cation influx. Ca2+
Photoreceptor depolarizes.

Photoreceptor
cell (rod)

Ca2+

Bipolar
cell

Ganglion
cell

© 2016 Pearson Education, Inc.


Figure 15.18-1 Signal transmission in the retina. Slide 3
Below, we look at a tiny column of retina.
The outer segment of the rod, closest to
the back of the eye and farthest from the
incoming light, is at the top.

In the dark

Light
1 cGMP-gated channels Na+
open, allowing cation influx. Ca2+
Photoreceptor depolarizes.

Photoreceptor
cell (rod)

2 Voltage-gated Ca2+
channels open in synaptic
terminals.
Ca2+

Bipolar
cell

Ganglion
cell

© 2016 Pearson Education, Inc.


Figure 15.18-1 Signal transmission in the retina. Slide 4
Below, we look at a tiny column of retina.
The outer segment of the rod, closest to
the back of the eye and farthest from the
incoming light, is at the top.

In the dark

Light
1 cGMP-gated channels Na+
open, allowing cation influx. Ca2+
Photoreceptor depolarizes.

Photoreceptor
cell (rod)

2 Voltage-gated Ca2+
channels open in synaptic
terminals.
Ca2+
3 Neurotransmitter is
released continuously.

Bipolar
cell

Ganglion
cell

© 2016 Pearson Education, Inc.


Figure 15.18-1 Signal transmission in the retina. Slide 5
Below, we look at a tiny column of retina.
The outer segment of the rod, closest to
the back of the eye and farthest from the
incoming light, is at the top.

In the dark

Light
1 cGMP-gated channels Na+
open, allowing cation influx. Ca2+
Photoreceptor depolarizes.

Photoreceptor
cell (rod)

2 Voltage-gated Ca2+
channels open in synaptic
terminals.
Ca2+
3 Neurotransmitter is
released continuously.

4 Neurotransmitter causes
IPSPs in bipolar cell.
Hyperpolarization results.

Bipolar
cell

Ganglion
cell

© 2016 Pearson Education, Inc.


Figure 15.18-1 Signal transmission in the retina. Slide 6
Below, we look at a tiny column of retina.
The outer segment of the rod, closest to
the back of the eye and farthest from the
incoming light, is at the top.

In the dark

Light
1 cGMP-gated channels Na+
open, allowing cation influx. Ca2+
Photoreceptor depolarizes.

Photoreceptor
cell (rod)

2 Voltage-gated Ca2+
channels open in synaptic
terminals.
Ca2+
3 Neurotransmitter is
released continuously.

4 Neurotransmitter causes
IPSPs in bipolar cell.
Hyperpolarization results.

Bipolar
cell
5 Hyperpolarization closes
voltage-gated Ca2+ channels,
inhibiting neurotransmitter
release.

Ganglion
cell

© 2016 Pearson Education, Inc.


Figure 15.18-1 Signal transmission in the retina. Slide 7
Below, we look at a tiny column of retina.
The outer segment of the rod, closest to
the back of the eye and farthest from the
incoming light, is at the top.

In the dark

Light
1 cGMP-gated channels Na+
open, allowing cation influx. Ca2+
Photoreceptor depolarizes.

Photoreceptor
cell (rod)

2 Voltage-gated Ca2+
channels open in synaptic
terminals.
Ca2+
3 Neurotransmitter is
released continuously.

4 Neurotransmitter causes
IPSPs in bipolar cell.
Hyperpolarization results.

Bipolar
cell
5 Hyperpolarization closes
voltage-gated Ca2+ channels,
inhibiting neurotransmitter
release.

6 No EPSPs occur in
ganglion cell. Ganglion
cell

© 2016 Pearson Education, Inc.


Figure 15.18-1 Signal transmission in the retina. Slide 8
Below, we look at a tiny column of retina.
The outer segment of the rod, closest to
the back of the eye and farthest from the
incoming light, is at the top.

In the dark

Light
1 cGMP-gated channels Na+
open, allowing cation influx. Ca2+
Photoreceptor depolarizes.

Photoreceptor
cell (rod)

2 Voltage-gated Ca2+
channels open in synaptic
terminals.
Ca2+
3 Neurotransmitter is
released continuously.

4 Neurotransmitter causes
IPSPs in bipolar cell.
Hyperpolarization results.

Bipolar
cell
5 Hyperpolarization closes
voltage-gated Ca2+ channels,
inhibiting neurotransmitter
release.

6 No EPSPs occur in
ganglion cell. Ganglion
cell
7 No action potentials occur
along the optic nerve.

© 2016 Pearson Education, Inc.


Figure 15.18-2 Signal transmission in the retina. Slide 2
Below, we look at a tiny column of retina.
The outer segment of the rod, closest to
the back of the eye and farthest from the
incoming light, is at the top.

In the light

Light
1 cGMP-gated channels
close, so cation influx
Light stops. Photoreceptor
hyperpolarizes.

Photoreceptor
cell (rod)

Bipolar
cell

Ca2+

Ganglion
cell

© 2016 Pearson Education, Inc.


Figure 15.18-2 Signal transmission in the retina. Slide 3
Below, we look at a tiny column of retina.
The outer segment of the rod, closest to
the back of the eye and farthest from the
incoming light, is at the top.

In the light

Light
1 cGMP-gated channels
close, so cation influx
Light stops. Photoreceptor
hyperpolarizes.

Photoreceptor
cell (rod)

2 Voltage-gated Ca2+
channels close in synaptic
terminals.

Bipolar
cell

Ca2+

Ganglion
cell

© 2016 Pearson Education, Inc.


Figure 15.18-2 Signal transmission in the retina. Slide 4
Below, we look at a tiny column of retina.
The outer segment of the rod, closest to
the back of the eye and farthest from the
incoming light, is at the top.

In the light

Light
1 cGMP-gated channels
close, so cation influx
Light stops. Photoreceptor
hyperpolarizes.

Photoreceptor
cell (rod)

2 Voltage-gated Ca2+
channels close in synaptic
terminals.

3 No neurotransmitter
is released.

Bipolar
cell

Ca2+

Ganglion
cell

© 2016 Pearson Education, Inc.


Figure 15.18-2 Signal transmission in the retina. Slide 5
Below, we look at a tiny column of retina.
The outer segment of the rod, closest to
the back of the eye and farthest from the
incoming light, is at the top.

In the light

Light
1 cGMP-gated channels
close, so cation influx
Light stops. Photoreceptor
hyperpolarizes.

Photoreceptor
cell (rod)

2 Voltage-gated Ca2+
channels close in synaptic
terminals.

3 No neurotransmitter
is released.

4 Lack of IPSPs in bipolar


cell results in depolarization.

Bipolar
cell

Ca2+

Ganglion
cell

© 2016 Pearson Education, Inc.


Figure 15.18-2 Signal transmission in the retina. Slide 6
Below, we look at a tiny column of retina.
The outer segment of the rod, closest to
the back of the eye and farthest from the
incoming light, is at the top.

In the light

Light
1 cGMP-gated channels
close, so cation influx
Light stops. Photoreceptor
hyperpolarizes.

Photoreceptor
cell (rod)

2 Voltage-gated Ca2+
channels close in synaptic
terminals.

3 No neurotransmitter
is released.

4 Lack of IPSPs in bipolar


cell results in depolarization.

Bipolar
cell
5 Depolarization opens
voltage-gated Ca2+ channels;
Ca2+ neurotransmitter is released.

Ganglion
cell

© 2016 Pearson Education, Inc.


Figure 15.18-2 Signal transmission in the retina. Slide 7
Below, we look at a tiny column of retina.
The outer segment of the rod, closest to
the back of the eye and farthest from the
incoming light, is at the top.

In the light

Light
1 cGMP-gated channels
close, so cation influx
Light stops. Photoreceptor
hyperpolarizes.

Photoreceptor
cell (rod)

2 Voltage-gated Ca2+
channels close in synaptic
terminals.

3 No neurotransmitter
is released.

4 Lack of IPSPs in bipolar


cell results in depolarization.

Bipolar
cell
5 Depolarization opens
voltage-gated Ca2+ channels;
Ca2+ neurotransmitter is released.

6 EPSPs occur in ganglion


cell.

Ganglion
cell

© 2016 Pearson Education, Inc.


Figure 15.18-2 Signal transmission in the retina. Slide 8
Below, we look at a tiny column of retina.
The outer segment of the rod, closest to
the back of the eye and farthest from the
incoming light, is at the top.

In the light

Light
1 cGMP-gated channels
close, so cation influx
Light stops. Photoreceptor
hyperpolarizes.

Photoreceptor
cell (rod)

2 Voltage-gated Ca2+
channels close in synaptic
terminals.

3 No neurotransmitter
is released.

4 Lack of IPSPs in bipolar


cell results in depolarization.

Bipolar
cell
5 Depolarization opens
voltage-gated Ca2+ channels;
Ca2+ neurotransmitter is released.

6 EPSPs occur in ganglion


cell.

7 Action potentials
Ganglion propagate along the
cell optic nerve.

© 2016 Pearson Education, Inc.


Light and Dark Adaptation

• Rhodopsin is so sensitive that bleaching


occurs even in starlight
• In bright light, bleaching occurs so fast that rods are
virtually nonfunctional
• Cones respond to bright light
• So, activation of rods and cones depends on:
• Light adaptation
• Dark adaptation

© 2016 Pearson Education, Inc.


Light and Dark Adaptation (cont.)

• Light adaptation
• When moving from darkness into bright light we see
glare because:
• Both rods and cones are strongly stimulated
• Large amounts of pigments are broken down
instantaneously, producing glare
• Pupils constrict
• Visual acuity improves over 5–10 minutes as:
• Rod system turns off
• Retinal sensitivity decreases
• Cones and neurons rapidly adapt

© 2016 Pearson Education, Inc.


Light and Dark Adaptation (cont.)

• Dark adaptation
• When moving from bright light into darkness, we
see blackness because:
• Cones stop functioning in low-intensity light
• Bright light bleached rod pigments, so they are still turned
off
• Pupils dilate
• Rhodopsin accumulates in dark, so retinal
sensitivity starts to increase
• Transducin returns to outer segments
• Sensitivity increases within 20–30 minutes

© 2016 Pearson Education, Inc.


Clinical – Homeostatic Imbalance 15.10

• Nyctalopia (night blindness): condition in which


rod function is seriously hampered
• Ability to drive safely at night is impaired
• Due to rod degeneration, commonly caused by
prolonged vitamin A deficiency
• If administered early, vitamin A supplements restore
function
• Can also be caused by retinitis pigmentosa
• Degenerative retinal diseases that destroy rods
• Tips of rods are not replaced when they slough off

© 2016 Pearson Education, Inc.


15.4 Processing and Relaying of Visual Information

Visual Pathway to the Brain


• Axons of retinal ganglion cells form optic
nerve
• Medial fibers from each eye cross over at the
optic chiasma then continue on as optic
tracts, which means each optic tract:
• Contains fibers from lateral (temporal) aspect of eye
on same side and medial (nasal) aspect of opposite
eye, and
• Each carries information from same half of visual
field
© 2016 Pearson Education, Inc.
Visual Pathway to the Brain (cont.)

• Most fibers of optic tracts continue on to lateral


geniculate nuclei of thalamus
• From there, thalamic neurons form optic
radiation, which projects to primary visual
cortex in occipital lobes
• Conscious perception of visual images occurs here

© 2016 Pearson Education, Inc.


Visual Pathway to the Brain (cont.)

• Other optic tract fibers send branches to


midbrain
• One set ends in superior colliculi, area controlling
extrinsic eye muscles
• A small subset of ganglion cells in retina
contains melanopsin (circadian pigment),
which projects to:
• Pretectal nuclei: involved with pupillary reflexes
• Suprachiasmatic nucleus of hypothalamus: timer
for daily biorhythms

© 2016 Pearson Education, Inc.


Figure 15.19a Visual pathway to the brain and visual fields, inferior view.

Fixation point

Right eye Left eye


Optic nerve
Supra-
chiasmatic
nucleus Optic chiasma

Pretectal Optic tract


nucleus

Uncrossed
Lateral (ipsilateral) fiber
geniculate
nucleus of Crossed
thalamus (contralateral) fiber

Optic
Superior
Primary visual radiation
colliculus
cortex
(occipital lobe)

The visual fields of the two eyes overlap considerably.


Note that fibers from the lateral portion of each retinal field
do not cross at the optic chiasma.
© 2016 Pearson Education, Inc.
Figure 15.19 Visual pathway to the brain and visual fields, inferior view.

Fixation point

Right eye Left eye


Optic nerve
Supra-
chiasmatic
nucleus Optic chiasma

Pretectal Optic tract


nucleus
Lateral
geniculate
nucleus
Superior
colliculus
(sectioned)

Uncrossed
Lateral (ipsilateral) fiber
geniculate
nucleus of Crossed
thalamus (contralateral) fiber

Optic
Superior
Primary visual radiation Corpus
colliculus
cortex callosum
(occipital lobe)

The visual fields of the two eyes overlap considerably. Photograph of human brain, with
Note that fibers from the lateral portion of each retinal field the right side dissected to reveal
do not cross at the optic chiasma. internal structures.
© 2016 Pearson Education, Inc.
Depth Perception

• Both eyes view same image from slightly


different angles
• Visual cortex fuses these slightly different
images, resulting in a three-dimensional image,
which leads to depth perception
• Requires input from both eyes

© 2016 Pearson Education, Inc.


Clinical – Homeostatic Imbalance 15.11

• Loss of an eye or destruction of one optic


nerve eliminates true depth perception entirely
• Peripheral vision on damaged side is also affected
• If neural destruction occurs beyond optic
chiasma, then part or all of opposite half of the
visual field is lost
• Example: stroke can affect left visual cortex,
which leads to blindness in right half of visual
field

© 2016 Pearson Education, Inc.


Visual Processing

• Retinal cells split input into channels that


include information about:
• Color and brightness, but also complex info such as
angle, direction, and speed of movement of edges
(sudden changes in brightness or color)
• Lateral inhibition decodes “edge” information
• Job of amacrine and horizontal cells

© 2016 Pearson Education, Inc.


Visual Processing (cont.)

• Ganglions pass information to lateral


geniculate nuclei of thalamus to be processed
for depth perception, with cone input
emphasized
• Primary visual cortex contains topographical
map of retina
• Neurons here respond to dark and bright edges and
to object orientation
• Provide form, color, motion inputs to visual
association areas

© 2016 Pearson Education, Inc.


Visual Processing (cont.)

• Info is also passed on to temporal, parietal,


and frontal lobes, where objects are identified
and location in space determined

© 2016 Pearson Education, Inc.


Part 2 – The Chemical Senses: Smell and Taste

• Smell (olfaction) and taste (gustation):


complementary senses that let us know
whether a substance should be savored or
avoided
• Chemoreceptors are used by these systems
• Chemicals must be dissolved in aqueous solution to
be picked up by chemoreceptors
• Smell receptors are excited by chemicals dissolved in
nasal fluids
• Taste receptors respond to chemicals dissolved in saliva

© 2016 Pearson Education, Inc.


15.5 Sense of Smell

Location and Structure of Olfactory


Receptors
• Olfactory epithelium: organ of smell
• Located in in roof of nasal cavity
• Covers superior nasal conchae
• Contains olfactory sensory neurons
• Bipolar neurons with radiating olfactory cilia
• Supporting cells surround and cushion olfactory receptor
cells
• Olfactory stem cells lie at base of epithelium

© 2016 Pearson Education, Inc.


Figure 15.20a Olfactory receptors.

Olfactory
epithelium

Olfactory tract

Olfactory bulb

Nasal
conchae

Route of
inhaled air

© 2016 Pearson Education, Inc.


Location and Structure of Olfactory Receptors (cont.)

• Olfactory neurons are unusual bipolar neurons


• Thin apical dendrites terminate in knob
• Long, largely nonmotile cilia, olfactory cilia, radiate
from knob
• Covered by mucus (solvent for odorants)
• Bundles of nonmyelinated axons of olfactory
receptor cells gather in fascicles that make up
filaments of olfactory nerve (cranial nerve I)
• Olfactory neurons, unlike other neurons, have
stem cells that give rise to new neurons every
30–60 days
© 2016 Pearson Education, Inc.
Figure 15.20b Olfactory receptors.

Mitral cell
Olfactory (output cell)
tract
Glomeruli

Olfactory bulb

Cribriform plate
of ethmoid bone

Filaments of
olfactory nerve
Lamina propria
Olfactory connective tissue
gland Olfactory axon
Olfactory stem cell
Olfactory sensory
Olfactory neuron
epithelium
Supporting cell
Dendrite

Mucus Olfactory cilia

Route of inhaled air


containing odor molecules

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Specificity of Olfactory Receptors

• Smells may contain 100s of different odorants


• Humans have ~400 “smell” genes active in
nose
• Each encodes a unique receptor protein
• Protein responds to one or more odors
• Each odor binds to several different receptors
• Each receptor has one type of receptor protein
• Pain and temperature receptors are also in
nasal cavities
• Respond to irritants, such as ammonia, or can
“smell” hot or cold (chili peppers, menthol)
© 2016 Pearson Education, Inc.
Physiology of Smell

• In order to smell substance, it must be volatile


• Must be in gaseous state
• Odorant must also be able to dissolve in olfactory
epithelium fluid
• Activation of olfactory sensory neurons
• Dissolved odorants bind to receptor proteins in
olfactory cilium membranes
• Open cation channels, generating receptor potential
• At threshold, AP is conducted to first relay station in
olfactory bulb

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Physiology of Smell (cont.)

• Smell transduction
• Odorant binds to receptor, activating a G protein
• Referred to as Golf
• G protein activation causes cAMP (second
messenger) synthesis
• cAMP opens Na+ and Ca2+ channels
• Na+ influx causes depolarization and impulse
transmission

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Physiology of Smell (cont.)

• Smell transduction (cont.)


• Ca2+ influx causes decreased response to a
sustained stimulus, referred to as olfactory
adaptation
• People can’t smell a certain odor after being exposed to it
for a while

© 2016 Pearson Education, Inc.


Figure 15.21 Olfactory transduction process. Slide 2

Odorant 1 Odorant binds Na+ Ca2+


to its receptor.

G protein (Golf)

cAMP
cAMP
Open cAMP-gated
GTP ATP cation channel
Receptor GTP

GDP GTP

© 2016 Pearson Education, Inc.


Figure 15.21 Olfactory transduction process. Slide 3

Odorant 1 Odorant binds Na+ Ca2+


to its receptor.

G protein (Golf)

cAMP
cAMP
Open cAMP-gated
GTP ATP cation channel
Receptor GTP

2 Receptor
GDP GTP activates G
protein (Golf).

© 2016 Pearson Education, Inc.


Figure 15.21 Olfactory transduction process. Slide 4

Odorant 1 Odorant binds Adenylate cyclase Na+ Ca2+


to its receptor.

G protein (Golf)

cAMP
cAMP
Open cAMP-gated
GTP ATP cation channel
Receptor GTP

2 Receptor 3 G protein
GDP GTP activates G activates adenylate
protein (Golf). cyclase.

© 2016 Pearson Education, Inc.


Figure 15.21 Olfactory transduction process. Slide 5

Odorant 1 Odorant binds Adenylate cyclase Na+ Ca2+


to its receptor.

G protein (Golf)

cAMP
cAMP
Open cAMP-gated
GTP ATP cation channel
Receptor GTP

2 Receptor 3 G protein 4 Adenylate cyclase


GDP GTP activates G activates adenylate converts ATP to cAMP.
protein (Golf). cyclase.

© 2016 Pearson Education, Inc.


Figure 15.21 Olfactory transduction process. Slide 6

Odorant 1 Odorant binds Adenylate cyclase Na+ Ca2+


to its receptor.

G protein (Golf)

cAMP
cAMP
Open cAMP-gated
GTP ATP cation channel
Receptor GTP

2 Receptor 3 G protein 4 Adenylate cyclase 5 cAMP opens a cation


GDP GTP activates G activates adenylate converts ATP to cAMP. channel, allowing Na+
protein (Golf). cyclase. and Ca2+ influx and
causing depolarization.

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The Olfactory Pathway

• Filaments of olfactory nerves synapse with


mitral cells located in overlying olfactory bulb
• Mitral cells are second-order neurons that form
olfactory tract
• Synapse occurs in structures called glomeruli
• Axons from neurons with same receptor type
converge on given type of glomerulus
• Mitral cells amplify, refine, and relay signals
• Amacrine granule cells release GABA to inhibit
mitral cells so that only highly excitatory
impulses are transmitted
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The Olfactory Pathway (cont.)

• Impulses from activated mitral cells travel via


olfactory tracts to piriform lobe of olfactory
cortex
• Some information sent to frontal lobe, and
some passes through thalamus first
• Smell is consciously interpreted and identified
• Some information sent to hypothalamus,
amygdala, and other regions of limbic system
• Emotional responses to odor are elicited

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Clinical – Homeostatic Imbalance 15.12

• Anosmias: olfactory disorders; most result


from
• Head injuries that tear olfactory nerves
• Aftereffects of nasal cavity inflammation
• Neurological disorders, such as Parkinson’s
disease
• Olfactory hallucinations
• Usually caused by temporal lobe epilepsy that
involves olfactory cortex
• Some people have olfactory auras prior to epileptic
seizures

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15.6 Sense of Taste

Location and Structure of Taste Buds


• Taste buds: sensory organs for taste
• Most of 10,000 taste buds are located on tongue in
papillae, peglike projections of tongue mucosa
• Fungiform papillae: tops of these mushroom-shaped
structures house most taste buds; scattered across
tongue
• Foliate papillae: on side walls of tongue
• Vallate papillae: largest taste buds with 8–12 forming “V”
at back of tongue
• Few on soft palate, cheeks, pharynx, epiglottis

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Figure 15.22a Location and structure of taste buds on the tongue.

Epiglottis

Palatine
tonsil

Lingual
tonsil

Foliate
papillae

Fungiform
papillae

Taste buds are associated


with fungiform, foliate,
and vallate papillae.
© 2016 Pearson Education, Inc.
Location and Structure of Taste Buds (cont.)

• Each taste bud consists of 50–100 flask-


shaped epithelial cells of two types:
• Gustatory epithelial cells: taste receptor cells
have microvilli called gustatory hairs that project
into taste pores, bathed in saliva
• Sensory dendrites coiled around gustatory epithelial cells
send taste signals to brain
• Three types of gustatory epithelial cells
• One releases serotonin; others lack synaptic vesicles, but
one releases ATP as neurotransmitter
• Basal epithelial cells: dynamic stem cells that
divide every 7–10 days

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Figure 15.22b Location and structure of taste buds on the tongue.

Vallate
papilla

Taste bud

Enlarged section of a
vallate papilla.

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Figure 15.22c Location and structure of taste buds on the tongue.

Connective
tissue Gustatory
hair
Taste fibers
of cranial
nerve

Stratified
Basal Gustatory Taste squamous
epithelial epithelial pore epithelium
cells cells of tongue

Enlarged view of a taste bud (210×).


© 2016 Pearson Education, Inc.
Basic Taste Sensations

• There are five basic taste sensations


1. Sweet—sugars, saccharin, alcohol, some
amino acids, some lead salts
2. Sour—hydrogen ions in solution
3. Salty—metal ions (inorganic salts); sodium
chloride tastes saltiest
4. Bitter—alkaloids such as quinine and nicotine,
caffeine, and nonalkaloids such as aspirin
5. Umami—amino acids glutamate and aspartate;
example: beef (meat) or cheese taste, and
monosodium glutamate

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Basic Taste Sensations (cont.)

• Possible sixth taste


• Growing evidence humans can taste long-chain
fatty acids from lipids
• Perhaps explain liking of fatty foods
• Taste likes/dislikes have homeostatic value
• Guide intake of beneficial and potentially harmful
substances
• Dislike for sourness and bitterness is a protective
way of warning us if something is spoiled or
poisonous

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Physiology of Taste

• To be able to taste a chemical, it must:


• Be dissolved in saliva
• Diffuse into taste pore
• Contact gustatory hairs

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Physiology of Taste (cont.)

• Activation of taste receptors


• Binding of food chemical (tastant) depolarizes cell
membrane of gustatory epithelial cell membrane,
causing release of neurotransmitter
• Neurotransmitter binds to dendrite of sensory neuron and
initiates a generator potential that lead to action potentials
• Different gustatory cells have different thresholds
for activation
• Bitter receptors are most sensitive
• All adapt in 3–5 seconds, with complete adaptation
in 1–5 minutes

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Physiology of Taste (cont.)

• Taste transduction
• Gustatory epithelial cell depolarization caused by:
• Salty taste is due to Na+ influx that directly causes
depolarization
• Sour taste is due to H+ acting intracellularly by opening
channels that allow other cations to enter
• Unique receptors for sweet, bitter, and umami, but all are
coupled to G protein gustducin
• Activation causes release of stored Ca2+ that opens cation
channels, causing depolarization and release of
neurotransmitter ATP

© 2016 Pearson Education, Inc.


Gustatory Pathway

• Two main cranial nerve pairs carry taste


impulses from tongue to brain:
• Facial nerve (VII) carries impulses from anterior
two-thirds of tongue
• Glossopharyngeal (X) carries impulses from
posterior one-third and pharynx
• Vagus nerve transmits from epiglottis and lower
pharynx

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Gustatory Pathway (cont.)

• Fibers synapse in the solitary nucleus of the


medulla, then travel to thalamus, and then to
gustatory cortex in the insula
• Hypothalamus and limbic system are involved;
allow us to determine appreciation of taste

© 2016 Pearson Education, Inc.


Figure 15.23 The gustatory pathway.

Gustatory
cortex
(in insula)

Thalamic
nucleus
(ventral
posteromedial
Pons
nucleus) Solitary nucleus
in medulla
oblongata
Facial
nerve (VII) Vagus nerve (X)
Glossopharyngeal
nerve (IX)

© 2016 Pearson Education, Inc.


Gustatory Pathway (cont.)

• Important roles of taste involve:


• Triggering reflexes involved in digestion, such as:
• Increased secretion of saliva into mouth
• Increased secretion of gastric juice into stomach
• May initiate protective reactions, such as:
• Gagging
• Reflexive vomiting

© 2016 Pearson Education, Inc.


Influence of Other Sensations on Taste

• Taste is 80% smell


• If nose is blocked, foods taste bland
• Mouth also contains thermoreceptors,
mechanoreceptors, and nociceptors
• Temperature and texture enhance or detract from
taste
• Spicy hot foods can excite pain receptors in mouth,
which some people experience as pleasure
• Example: hot chili peppers

© 2016 Pearson Education, Inc.


Clinical – Homeostatic Imbalance 15.13

• Taste disorders are less common than


disorders of smell, mostly because taste
receptors are served by three different nerves
• Not likely that all three nerves would be damaged at
same time

© 2016 Pearson Education, Inc.


Clinical – Homeostatic Imbalance 15.13

• Causes of taste disorders include:


• Upper respiratory tract infections
• Head injuries
• Chemicals or medications
• Head and neck radiation for cancer treatment
• Zinc supplements may help some cases of radiation-
induced taste disorders

© 2016 Pearson Education, Inc.


Part 3 – The Ear: Hearing and Balance

15.7 Structure of the Ear

• The ear has three major areas:


• External (outer) ear: hearing only
• Middle ear (tympanic cavity): hearing only
• Internal (inner) ear: hearing and equilibrium
• Receptors for hearing and balance respond to
separate stimuli and are activated
independently

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External Ear

• The external (outer) ear consists of two parts:


• Auricle (pinna): shell-shaped structure surrounding
ear canal that functions to funnel sound waves into
auditory canal
• Helix: cartilaginous rim
• Lobule: fleshy earlobe
• External acoustic meatus (auditory canal)
• Short, curved tube lined with skin bearing hairs,
sebaceous glands, and ceruminous (earwax) glands
• Transmits sound waves to eardrum

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External Ear (cont.)

• Tympanic membrane (eardrum)


• Boundary between external and middle ears
• Thin, translucent connective tissue membrane
• Vibrates in response to sound
• Transfers sound energy to bones of middle ear

© 2016 Pearson Education, Inc.


Figure 15.24a Structure of the ear.

External Middle Internal ear


ear ear (labyrinth)

Auricle
(pinna)

Helix

Lobule
External
acoustic
Tympanic Pharyngotympanic
meatus
membrane (auditory) tube
The three regions of the ear
© 2016 Pearson Education, Inc.
Middle Ear (Tympanic Cavity)

• A small, air-filled, mucosa-lined cavity in


temporal bone
• Flanked laterally by eardrum and medially by bony
wall containing oval and round membranous
windows
• Epitympanic recess: superior portion of
middle ear (roof of cavity)
• Mastoid antrum: canal for communication with
mastoid air cells in mastoid process

© 2016 Pearson Education, Inc.


Figure 15.24b Structure of the ear.

Oval window
(deep to stapes)
Entrance to mastoid Semicircular
antrum in the canals
epitympanic recess
Malleus Vestibule
(hammer)

Auditory Incus Vestibular


ossicles (anvil) nerve
Stapes
Cochlear
(stirrup)
nerve
Tympanic
membrane Cochlea

Round window Pharyngotympanic


(auditory) tube

Middle and internal ear

© 2016 Pearson Education, Inc.


Middle Ear (Tympanic Cavity) (cont.)

• Pharyngotympanic (auditory) tube: connects


middle ear to nasopharynx
• Formerly called eustachian tube
• Usually flattened tube, but can be opened by
yawning or swallowing to equalize pressure in
middle ear cavity with external air pressure
• Tympanic membrane cannot vibrate efficiently if pressures
on both sides are not equal
• Sounds are distorted

© 2016 Pearson Education, Inc.


Middle Ear (Tympanic Cavity) (cont.)

• Auditory ossicles: three small bones in


tympanic cavity, named for their shape:
• Malleus: the “hammer” is secured to eardrum
• Incus: the “anvil”
• Stapes: the “stirrup” base fits into oval window
• Synovial joints allow malleus to articulate with
incus, which articulates with stapes
• Suspended by ligaments; transmit vibratory motion
of eardrum to oval window
• Tensor tympani and stapedius muscles contract
reflexively in response to loud sounds to prevent
© 2016 Pearson Education, Inc.
Figure 15.25 The three auditory ossicles and associated skeletal muscles.

View
Epitympanic
Malleus Incus recess
Superior

Lateral

Anterior

Pharyngotym- Tensor Tympanic Stapes Stapedius


panic tube tympani membrane muscle
muscle (medial view)
© 2016 Pearson Education, Inc.
Clinical – Homeostatic Imbalance 15.14

• Otitis media
• Middle ear inflammation
• Commonly seen in children with sore throat
• Especially those with shorter, more horizontal
pharyngotympanic tubes
• Most frequent cause of hearing loss in children
• Acute infectious forms cause eardrum to bulge
outward and become inflamed
• Most cases respond to antibiotics

© 2016 Pearson Education, Inc.


Internal Ear

• Also referred to as the labyrinth (maze)


• Located in temporal bone behind eye socket
• Two major divisions:
• Bony labyrinth: system of tortuous channels and
cavities that worm through the bone
• Divided into three regions: vestibule, semicircular
canals, and cochlea
• Filled with perilymph fluid; similar to CSF
• Membranous labyrinth; series of membranous
sacs and ducts contained in bony labyrinth; filled
with potassium-rich endolymph

© 2016 Pearson Education, Inc.


Figure 15.26 Membranous labyrinth of the internal ear.

Temporal
bone

Facial nerve
Semicircular ducts in
semicircular canals Vestibular nerve
• Anterior
• Posterior Superior vestibular ganglion
• Lateral Inferior vestibular ganglion

Cristae ampullares Cochlear nerve


in the membranous Maculae
ampullae
Spiral organ
Utricle in vestibule
Cochlear duct
Saccule in vestibule in cochlea
Stapes in
oval window Round window

© 2016 Pearson Education, Inc.


Internal Ear (cont.)

• Vestibule
• Central egg-shaped cavity of bony labyrinth
• Contains two membranous sacs
1. Saccule is continuous with cochlear duct
2. Utricle is continuous with semicircular canals
• Sacs house equilibrium receptor regions (maculae)
that respond to gravity and changes in position of
head

© 2016 Pearson Education, Inc.


Internal Ear (cont.)

• Semicircular canals
• Three canals oriented in three planes of space:
anterior, lateral, and posterior
• Anterior and posterior are at right angles to each other,
whereas the lateral canal is horizontal
• Membranous semicircular ducts line each canal
and communicate with utricle
• Ampulla: enlarged area of ducts of each canal that
houses equilibrium receptor region called the crista
ampullaris
• Receptors respond to angular (rotational) movements of
the head

© 2016 Pearson Education, Inc.


Figure 15.26 Membranous labyrinth of the internal ear.

Temporal
bone

Facial nerve
Semicircular ducts in
semicircular canals Vestibular nerve
• Anterior
• Posterior Superior vestibular ganglion
• Lateral Inferior vestibular ganglion

Cristae ampullares Cochlear nerve


in the membranous Maculae
ampullae
Spiral organ
Utricle in vestibule
Cochlear duct
Saccule in vestibule in cochlea
Stapes in
oval window Round window

© 2016 Pearson Education, Inc.


Internal Ear (cont.)

• Cochlea
• A small spiral, conical, bony chamber, size of a split
pea
• Extends from vestibule
• Coils around bony pillar (modiolus)
• Contains cochlear duct, which houses spiral organ
(organ of Corti) and ends at cochlear apex

© 2016 Pearson Education, Inc.


Internal Ear (cont.)

• Cochlea (cont.)
• Cavity of cochlea divided into three chambers:
• Scala vestibule: abuts oval window, contains
perilymph
• Scala media (cochlear duct): contains endolymph
• Scala tympani: terminates at round window;
contains perilymph

© 2016 Pearson Education, Inc.


Internal Ear (cont.)

• Cochlea (cont.)
• Scalae tympani and vestibuli are continuous
with each other at helicotrema (apex)
• Vestibular membrane: “roof” of cochlear duct
that separates scala media from scala vestibuli

© 2016 Pearson Education, Inc.


Internal Ear (cont.)

• Cochlea (cont.)
• Stria vascularis: external wall of cochlear duct
composed of mucosa that secretes endolymph
• “Floor” of cochlear duct composed of:
• Bony spiral lamina
• Basilar membrane, which supports spiral organ

© 2016 Pearson Education, Inc.


Internal Ear (cont.)

• Cochlea (cont.)
• Spiral organ contains cochlear hair cells
functionally arranged in one row of inner hair
cells and three rows of outer hair cells
• Hair cells are sandwiched between tectorial and
basilar membranes
• The cochlear branch of nerve VIII runs from
spiral organ to brain

© 2016 Pearson Education, Inc.


Figure 15.27a Anatomy of the cochlea.

Helicotrema Modiolus
at apex

Cochlear nerve,
division of the
vestibulocochlear
nerve (VIII)

Spiral ganglion
Osseous spiral lamina

Vestibular membrane

Cochlear duct
(scala media)

© 2016 Pearson Education, Inc.


Figure 15.27b Anatomy of the cochlea.

Vestibular membrane Osseous spiral lamina


Tectorial membrane
Scala Spiral
Cochlear duct vestibuli ganglion
(scala media; (contains
contains perilymph)
endolymph)
Stria
vascularis

Spiral organ Scala


tympani
Basilar (contains
membrane perilymph)

© 2016 Pearson Education, Inc.


Figure 15.27c Anatomy of the cochlea.

Tectorial membrane Inner hair cell

Hairs (stereocilia) Afferent nerve


fibers
Outer hair cells

Supporting cells

Fibers of
cochlear
nerve

Basilar
membrane

© 2016 Pearson Education, Inc.


Figure 15.27d Anatomy of the cochlea.

Hairs
of inner
hair cell

Hairs
of outer
hair cell

© 2016 Pearson Education, Inc.


Table 15.2 Summary of the Internal Ear

© 2016 Pearson Education, Inc.


Hearing

15.8 Sound Detection

• Hearing is the reception of an air sound wave


that is converted to a fluid wave that ultimately
stimulates mechanosensitive cochlear hair
cells that send impulses to the brain for
interpretation
Properties of Sound
• Sound is a pressure disturbance (alternating
areas of high and low pressure) produced by a
vibrating object and propagated by molecules
of the medium (air)

© 2016 Pearson Education, Inc.


Properties of Sound (cont.)

• Sound waves are created when an object


moves:
• Air molecules that are displaced by object
movement are pushed forward into adjacent area,
adding to air molecules already there
• Creates an area of high pressure due to compression of
molecules together
• As object returns to original position, the area it
leaves now has fewer air molecules
• Creates an area of low pressure due to presence of fewer
air molecules
• Referred to as rarefaction

© 2016 Pearson Education, Inc.


Properties of Sound (cont.)

• Sound waves are alternating areas (waves) of


compressions and rarefactions
• Object vibrating causes waves to move
outward in all directions as air all around it is
compressed and rarefied
• Kinetic energy of object is transferred to air
molecules, which then transfer it to other air
molecules
• Wave energy declines with time and distance

© 2016 Pearson Education, Inc.


Properties of Sound (cont.)

• Illustrated as an S-shaped curve, or sine wave


• Compressions shown as crests, rarefactions as
troughs

© 2016 Pearson Education, Inc.


Figure 15.28a Sound: Source and propagation.

Area of
high pressure
(compressed
molecules)

Area of
Wavelength low pressure
(rarefaction)
Air pressure

Crest

Trough

Distance Amplitude
A struck tuning fork alternately compresses
and rarefies the air molecules around it.

© 2016 Pearson Education, Inc.


Figure 15.28b Sound: Source and propagation.

Sound waves
radiate outward
© 2016 Pearson Education, Inc.
in all directions.
Properties of Sound (cont.)

• Sound can be described by two physical


properties: frequency and amplitude
1. Frequency
• Number of waves that pass given point in a given time
• Pure tone has repeating crests and troughs
• Wavelength
• Distance between two consecutive crests
• Shorter wavelength = higher frequency of sound
• Wavelength is consistent for a particular sound

© 2016 Pearson Education, Inc.


Properties of Sound (cont.)

1. Frequency (cont.)
• Frequency range of human hearing is 20–20,000 hertz
(Hz = waves per second), but most sensitive between
1500 and 4000 Hz
• Pitch: perception of different frequencies
• Higher the frequency, higher the pitch
• Quality: characteristic of sounds
• Most sounds are mixtures of different frequencies
• Tone: one frequency (ex: tuning fork)
• Sound quality provides richness and complexity of
sounds (music)

© 2016 Pearson Education, Inc.


Figure 15.29a Frequency and amplitude of sound waves.

Pressure

High frequency
(short wavelength)
= high pitch

Low frequency
(long wavelength)
= low pitch
0.01 0.02 0.03
Time (s)
Frequency is perceived as pitch.

© 2016 Pearson Education, Inc.


Properties of Sound (cont.)

2. Amplitude
• Height of crests
• Amplitude perceived as loudness: subjective
interpretation of sound intensity
• Measured in decibels (dB)
• Normal range is 0–120 decibels (dB)
• Normal conversation is around 50 dB
• Threshold of pain is 120 dB
• Severe hearing loss can occur with prolonged exposure
above 90 dB
• Amplified rock music is 120 dB or more

© 2016 Pearson Education, Inc.


Figure 15.29b Frequency and amplitude of sound waves.

Pressure

High amplitude
= loud

Low amplitude
= soft
0.01 0.02 0.03
Time (s)
Amplitude (size or intensity) is perceived as loudness.

© 2016 Pearson Education, Inc.


Transmission of Sound to Internal Ear

• Pathway of sound
1. Tympanic membrane: sound waves enter
external acoustic meatus and strike tympanic
membrane, causing it to vibrate
• The higher the intensity, the more vibration
2. Auditory ossicles: transfer vibration of
eardrum to oval window
• Tympanic membrane is about 20 larger than oval
window, so vibration transferred to oval window is
amplified about 20

© 2016 Pearson Education, Inc.


Transmission of Sound to Internal Ear (cont.)

3. Scala vestibuli: stapes rocks back and forth


on oval window with each vibration, causing
wave motions in perilymph
• Wave ends at round window, causing it to bulge outward into
middle ear cavity
4a. Helicotrema path: waves with frequencies below
threshold of hearing travel through helicotrema and
scali tympani to round window
4b. Basilar membrane path: sounds in hearing range go
through cochlear duct, vibrating basilar membrane at
specific location, according to frequency of sound

© 2016 Pearson Education, Inc.


Figure 15.30 Pathway of sound waves. Slide 2

Auditory ossicles Cochlear


Malleus Incus Stapes
nerve 1 Sound waves vibrate the
tympanic membrane.
Oval Scala vestibuli
window
Helicotrema Scala
tympani
Cochlear
duct
Basilar
membrane
1

Tympanic Round
membrane window

© 2016 Pearson Education, Inc.


Figure 15.30 Pathway of sound waves. Slide 3

Auditory ossicles Cochlear


Malleus Incus Stapes
nerve 1 Sound waves vibrate the
tympanic membrane.
Oval Scala vestibuli
window 2 Auditory ossicles vibrate.
Helicotrema Scala Pressure is amplified.
tympani
Cochlear
2 duct
Basilar
membrane
1

Tympanic Round
membrane window

© 2016 Pearson Education, Inc.


Figure 15.30 Pathway of sound waves. Slide 4

Auditory ossicles Cochlear


Malleus Incus Stapes
nerve 1 Sound waves vibrate the
tympanic membrane.
Oval Scala vestibuli
window 2 Auditory ossicles vibrate.
Helicotrema Scala Pressure is amplified.
tympani
3 Pressure waves created by the
Cochlear stapes pushing on the oval window
2 3 duct move through fluid in the scala
vestibuli.
Basilar
membrane
1

Tympanic Round
membrane window

© 2016 Pearson Education, Inc.


Figure 15.30 Pathway of sound waves. Slide 5

Auditory ossicles Cochlear


Malleus Incus Stapes
nerve 1 Sound waves vibrate the
tympanic membrane.
Oval Scala vestibuli
window 2 Auditory ossicles vibrate.
Helicotrema Scala Pressure is amplified.
4a tympani
3 Pressure waves created by the
Cochlear stapes pushing on the oval window
2 3 duct move through fluid in the scala
vestibuli.
Basilar
membrane 4a Sounds with frequencies below
1 hearing travel through the
helicotrema and do not excite hair
cells.

Tympanic Round
membrane window

© 2016 Pearson Education, Inc.


Figure 15.30 Pathway of sound waves. Slide 6

Auditory ossicles Cochlear


Malleus Incus Stapes
nerve 1 Sound waves vibrate the
tympanic membrane.
Oval Scala vestibuli
window 2 Auditory ossicles vibrate.
Helicotrema Scala Pressure is amplified.
4a tympani
3 Pressure waves created by the
Cochlear stapes pushing on the oval window
2 3 duct move through fluid in the scala
vestibuli.
4b Basilar
membrane 4a Sounds with frequencies below
1 hearing travel through the
helicotrema and do not excite hair
cells.
4b Sounds in the hearing range go
through the cochlear duct, vibrating
the basilar membrane and
deflecting hairs on inner hair cells.
Tympanic Round
membrane window

© 2016 Pearson Education, Inc.


Resonance of the Basilar Membrane

• Resonance: movement of different areas of


basilar membrane in response to a particular
frequency
• Basilar membrane changes along its length:
• Fibers near oval window are short and stiff
• Resonate with high-frequency waves
• Fibers near cochlear apex are longer, floppier
• Resonate with lower-frequency waves
• So basilar membrane mechanically processes
sound even before signals reach receptors

© 2016 Pearson Education, Inc.


Figure 15.31-1 Basilar membrane function.

Let’s uncoil the cochlea to see how it


separates different frequencies of sound
so that we can hear different pitches.
Stapes
Basilar membrane

© 2016 Pearson Education, Inc.


Figure 15.31-2 Basilar membrane function.

The properties of the basilar membrane change along its length.

Short, stiff fibers Long, floppy fibers

Base

As a result, different frequencies vibrate the basilar Apex


membrane in different places.

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Sound Transduction

• Excitation of inner hair cells


• Movement of basilar membrane deflects hairs of
inner hair cells
• Cochlear hair cells have microvilli that contain many
stereocilia (hairs) that bend at their base
• Longest hair cells are connected to shortest hair cells via
tip links
• Tip links, when pulled on, open ion channels they are
connected to
• Stereocilia project into K+-rich endolymph, with
longest hairs enmeshed in gel-like tectorial
membrane

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Figure 15.27c Anatomy of the cochlea.

Tectorial membrane Inner hair cell

Hairs (stereocilia) Afferent nerve


fibers
Outer hair cells

Supporting cells

Fibers of
cochlear
nerve

Basilar
membrane

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Sound Transduction (cont.)

• Excitation of inner hair cells (cont.)


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

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Figure 15.32 Bending of stereocilia opens or closes mechanically gated ion channels in hair cells.

Basilar membrane at rest Hairs bent toward tallest stereocilium Hairs bent away from tallest stereocilium

Tectorial
membrane K+, Ca2+ 1 Tip links tighten, 1 Tip links loosen,
A few
opening closing mechanically
Tip link channels
mechanically gated ion channels.
are open;
Stereocilia cell slightly
gated ion channels. 2 No cations enter;
depolarized 2 More cations cell hyperpolarizes.
enter; cell
depolarizes.
Hair cell

3 Neurotransmitter 3 Neurotransmitter
release. release.
4 Action potentials 4 Action potentials
Basilar
in cochlear nerve. in cochlear nerve.
membrane

Cochlear nerve axon

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Sound Transduction (cont.)

• Role of outer hair cells


• Nerve fibers coiled around hair cells of outer row
are efferent neurons that convey messages from
brain to ear
• Outer hair cells can contract and stretch, which
changes stiffness of basilar membrane
• This ability serves two functions:
• Increase “fine-tuning” responsiveness of inner hair cells by
amplifying motion of basilar membrane
• Protect inner hair cells from loud noises by decreasing
motion of basilar membrane

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15.9 Auditory Pathways to Brain

Auditory Pathway
• Neural impulses from cochlear bipolar cells
reach auditory cortex via following pathway:
• Spiral ganglion 
• Cochlear nuclei (medulla) 
• Superior olivary nucleus (pons-medulla) 
• Lateral lemniscus (tract) 
• Inferior colliculus (midbrain auditory reflex center

• Medial geniculate nucleus (thalamus) 
• Primary auditory cortex
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Auditory Pathway (cont.)

• Some fibers cross over, some do not; so both


auditory cortices receive input from both ears

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Figure 15.33 The auditory pathway.

Medial geniculate
nucleus of thalamus

Primary auditory
cortex in temporal lobe

Inferior colliculus

Lateral lemniscus

Superior olivary
nucleus (pons- Midbrain
medulla junction)

Cochlear nuclei

Medulla
Vibrations
Vestibulocochlear
nerve
Vibrations
Spiral ganglion
of cochlear nerve

Bipolar cell

Spiral organ
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Auditory Processing

• Perception of pitch: impulses from hair cells


in different positions along basilar membrane
are interpreted by brain as specific pitches
• Detection of loudness is determined by brain
as an increase in the number of action
potentials (frequency) that result when hair
cells experience larger deflections

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Auditory Processing (cont.)

• Localization of sound depends on relative


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

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Equilibrium

15.10 Maintenance of Equilibrium

• Equilibrium is response to various movements


of head that rely on input from inner ear, eyes,
and stretch receptors
• Vestibular apparatus: equilibrium receptors in
semicircular canals and vestibule
• Vestibular receptors monitor static equilibrium
• Semicircular canal receptors monitor dynamic
equilibrium

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Figure 15.24b Structure of the ear.

Oval window
(deep to stapes)
Entrance to mastoid Semicircular
antrum in the canals
epitympanic recess
Malleus Vestibule
(hammer)

Auditory Incus Vestibular


ossicles (anvil) nerve
Stapes
Cochlear
(stirrup)
nerve
Tympanic
membrane Cochlea

Round window Pharyngotympanic


(auditory) tube

Middle and internal ear

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The Maculae

• Maculae: sensory receptor organs that monitor


static equilibrium
• One organ located in each saccule wall and one in
each utricle wall
• Monitor the position of head in space
• Play a key role in control of posture
• Respond to linear acceleration forces, but not
rotation

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Figure 15.26 Membranous labyrinth of the internal ear.

Temporal
bone

Facial nerve
Semicircular ducts in
semicircular canals Vestibular nerve
• Anterior
• Posterior Superior vestibular ganglion
• Lateral Inferior vestibular ganglion

Cristae ampullares Cochlear nerve


in the membranous Maculae
ampullae
Spiral organ
Utricle in vestibule
Cochlear duct
Saccule in vestibule in cochlea
Stapes in
oval window Round window

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The Maculae (cont.)

• Anatomy of a macula
• Each is a flat epithelium patch containing hair cells
with supporting cells
• Hair cells have stereocilia and special “true
stereocilium” called kinocilium
• Located next to tallest stereocilia
• Stereocilia are embedded in otolith membrane,
jelly-like mass studded with otoliths (tiny CaCO3
stones)
• Otoliths increase membrane’s weight and increase its
inertia (resistance to changes in motion)

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The Maculae (cont.)

• Anatomy of a macula (cont.)


• Utricle maculae are horizontal with vertical hairs
• Respond to change along a horizontal plane, such as
tilting head
• Forward/backward movements stimulate utricle
• Saccule maculae are vertical with horizontal hairs
• Respond to change along a vertical plane
• Up/down movements stimulate saccule (Example:
acceleration of an elevator)

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The Maculae (cont.)

• Anatomy of a macula (cont.)


• Hair cells synapse with fibers of vestibular nerve
whose cell bodies are located in superior and
inferior vestibular ganglia
• Part of vestibulocochlear cranial nerve (VIII)

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Figure 15.34a Structure and function of a macula.
Macula of
utricle
Macula of
saccule

Kinocilium Otoliths Otolith


Stereocilia membrane

Hair
cells

Supporting
cells
Vestibular
nerve fibers
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The Maculae (cont.)

• Activating receptors of a macula


• Hair cells release neurotransmitter continuously
• Acceleration/deceleration causes a change in amount of
neurotransmitter released
• Leads to change in AP frequency to brain
• Density of otolith membrane causes it to lag behind
movement of hair cells when head changes
positions
• Base of stereocilia moves at same rate as head, but tips
embedded in otolith are pulled by lagging membrane,
causing hair to bend
• Ion channels open, and depolarization occurs

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The Maculae (cont.)

• Activating receptors of a macula (cont.)


• Bending of hairs in direction of kinocilia:
• Depolarizes hair cells
• Increases amount of neurotransmitter release
• More impulses travel up vestibular nerve to brain
• Bending of hairs away from kinocilia:
• Hyperpolarizes receptors
• Less neurotransmitter released
• Reduces rate of impulse generation
• Thus brain is informed of changing position of head

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Figure 15.34b Structure and function of a macula.
Head Upright
Gravity

Steady stream of
action potentials in
vestibular nerve

Head tilted forward


Force
• Hairs bend toward
kinocilium
• Hair cell depolarizes
• Nerve fiber excited
• Action potentials
in vestibular nerve

Force Head tilted backwards


• Hairs bend away from
kinocilium
• Hair cell hyperpolarizes
• Nerve fiber inhibited
• Action potentials in
vestibular nerve

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The Cristae Ampullares

• Receptor for rotational acceleration is crista


ampullaris (crista)
• Small elevation in ampulla of each semicircular
canal
• Cristae are excited by acceleration and
deceleration of head
• Major stimuli are rotational (angular) movements,
such as twirling of the body
• Semicircular canals are located in all three planes
of space, so cristae can pick up on all rotational
movements of head

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The Cristae Ampullares (cont.)

• Anatomy of a crista ampullaris


• Each crista has supporting cells and hair cells that
extend into gel-like mass called ampullary cupula
• Dendrites of vestibular nerve fibers encircle base of
hair cells
• Activating receptors of crista ampullaris
• Cristae respond to changes in velocity of rotational
movements of head
• Inertia in ampullary cupula causes endolymph in
semicircular ducts to move in direction opposite
body’s rotation, causing hair cells to bend
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Figure 15.35a Location, structure, and function of a crista ampullaris in the internal ear.

Ampullary cupula

Crista
ampullaris
Endolymph

Hair bundle
(kinocilium
plus stereocilia)

Crista Hair cell


ampullaris
Membranous
labyrinth
Supporting
Fibers of vestibular nerve
cell

Anatomy of a crista ampullaris in a semicircular canal

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Figure 15.35b Location, structure, and function of a crista ampullaris in the internal ear.

Ampullary cupula

Scanning electron micrograph


of a crista ampullaris (200×)

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The Cristae Ampullares (cont.)

• Activating receptors of crista ampullaris


(cont.)
• Bending hairs in cristae causes depolarization
• Rapid impulses reach brain at faster rate
• Bending of hairs in opposite direction causes
hyperpolarizations
• Fewer impulses reach brain
• Thus brain is informed of head rotations

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The Cristae Ampullares (cont.)

• Activating receptors of crista ampullaris


(cont.)
• Axes of hair cells in complementary semicircular
ducts are opposite
• Depolarization occurs in one ear, while hyperpolarization
occurs in other ear
• Endolymph will come to rest after a while, so this
system detects only changes in movements

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Figure 15.35c Location, structure, and function of a crista ampullaris in the internal ear.

Section of Ampullary cupula Fibers of Flow of endolymph


ampulla, vestibular
filled with nerve
endolymph

At rest, the cupula stands upright. During rotational acceleration, endolymph As rotational movement slows,
moves inside the semicircular canals in the endolymph keeps moving in the direction
direction opposite the rotation (it lags of rotation. Endolymph flow bends the
behind due to inertia). Endolymph flow cupula in the opposite direction from
bends the cupula and excites the hair cells. acceleration and inhibits the hair cells.
Movement of the ampullary cupula during rotational acceleration and deceleration

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The Cristae Ampullares (cont.)

• Vestibular nystagmus
• Semicircular canal impulses are linked to reflex
movements of eyes
• Nystagmus is strange eye movements during and
immediately after rotation
• Often accompanied by vertigo
• As rotation begins, eyes drift in direction opposite to
rotation; then CNS compensation causes rapid
jump toward direction of rotation
• As rotation ends, eyes continue in direction of spin,
then jerk rapidly in opposite direction

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Equilibrium Pathway to the Brain

• Equilibrium information goes to reflex centers


in brain stem
• Allows fast, reflexive responses to imbalance so we
don’t fall down
• Impulses from activated vestibular receptors
travel to either vestibular nuclei in brain stem or
to cerebellum
• Three modes of input for balance and
orientation:
• Vestibular receptors
• Visual receptors
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Figure 15.36 Neural pathways of the balance and orientation system.

Input: Information about the body’s position in space


comes from three main sources and is fed into two
major processing areas in the central nervous system.

Somatic receptors
Vestibular Visual
(skin, muscle
receptors receptors
and joints)

Vestibular
Cerebellum nuclei
(brain stem)

Central nervous
system processing

Oculomotor control Spinal motor control


(cranial nerve nuclei (cranial nerve XI nuclei
III, IV, VI) and vestibulospinal tracts)

(eye movements) (neck, limb, and trunk


movements)

Output: Responses by the central nervous system provide fast reflexive


control of the muscles serving the eyes, neck, limbs, and trunk.
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Clinical – Homeostatic Imbalance 15.15

• Equilibrium problems are usually unpleasant


and can cause nausea, dizziness, and loss of
balance
• Nystagmus in the absence of rotational stimuli
may be present

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Clinical – Homeostatic Imbalance 15.15

• Motion sickness: sensory inputs are


mismatched
• Visual input differs from equilibrium input
• Conflicting information causes motion sickness
• Warning signs are excess salivation, pallor, rapid
deep breathing, profuse sweating
• Treatment with antimotion drugs that depress
vestibular input, such as meclizine and
scopolamine

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15.11 Homeostatic Imbalances of Hearing

Deafness
• Conduction deafness
• Blocked sound conduction to fluids of internal ear
• Causes include impacted earwax, perforated eardrum,
otitis media, otosclerosis of the ossicles
• Sensorineural deafness
• Damage to neural structures at any point from
cochlear hair cells to auditory cortical cells
• Typically from gradual hair cell loss

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Deafness (cont.)

• Sensorineural deafness research is under way


to prod supporting cells to differentiate into hair
cells
• Cochlear implants that convert sound energy
into electrical signals are effective for
congenital or age/noise cochlear damage
• Inserted into drilled recess in temporal bone
• So effective that deaf children can learn to speak

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Figure 15.37 Boy with a cochlear implant.

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Tinnitus

• Ringing, buzzing, or clicking sound in ears in


absence of auditory stimuli
• Due to cochlear nerve degeneration,
inflammation of middle or internal ears, side
effects of aspirin

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Ménière’s Syndrome

• Labyrinth disorder that affects cochlea and


semicircular canals
• Causes vertigo, nausea, and vomiting
• Treatment: anti–motion sickness drugs in mild
cases or surgical removal of labyrinth in severe
cases

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Developmental Aspects of the Special
Senses

Taste and Smell


• All special senses are functional at birth
• Chemical senses: few problems occur until
fourth decade, when these senses begin to
decline
• Odor and taste detection is poor after 65

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Developmental Aspects of the Special
Senses

Vision
• Optic vesicles protrude from diencephalon
during week 4 of development
• Vesicles indent to form optic cups
• Stalks form optic nerves
• Later, lens forms from ectoderm
• Vision is not fully functional at birth; babies are
hyperopic because eyes are shortened
• See only gray tones
• Eye movements are uncoordinated
• Tearless for about 2 weeks
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Developmental Aspects of the Special
Senses

Vision (cont.)
• By 5 months of age, infants can follow objects,
but acuity is still poor
• Depth perception and color vision develop by
age 3
• Adult eye size reached around 8–9 years of
age
• Around year 40, lenses start to lose elasticity,
resulting in presbyopia

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Developmental Aspects of the Special
Senses

Vision (cont.)
• With age, lens loses clarity, dilator muscles are
less efficient; visual acuity is drastically
decreased by age 70
• Lacrimal glands less active, so eyes are dry, more
prone to infection

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Developmental Aspects of the Special
Senses

Hearing and Balance


• Ear development begins in 3-week embryo
• Inner ears develop from thickening of ectoderm
called otic placodes, which invaginate into
otic pit and otic vesicle
• Otic vesicle becomes membranous labyrinth,
and surrounding mesenchyme becomes bony
labyrinth

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Developmental Aspects of the Special
Senses

Hearing and Balance (cont.)


• Middle ear structures develop from endodermal
pharyngeal pouches, ossicles from neural
crest cells, and pharyngeal cleft (branchial
groove) develops into outer ear structures
• Newborns can hear, but early responses are
reflexive in nature
• By month 4, infants can turn head toward voices of
family members

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Developmental Aspects of the Special
Senses

Hearing and Balance (cont.)


• Language skills tied to ability to hear well
• Few ear problems until 60s, when deterioration
of spiral organ becomes noticeable
• Hair cell numbers decline with age
• Presbycusis: loss of high-pitch perception occurs
first
• Type of sensorineural deafness

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Clinical – Homeostatic Imbalance 15.16

• Congenital problems of eyes are relatively


uncommon, but incidence is increased by
certain maternal infections
• Rubella (German measles) is dangerous,
especially during critical first 3 months of
pregnancy
• Common problems associated with rubella
infections are blindness and cataracts

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Clinical – Homeostatic Imbalance 15.17

• Congenital abnormalities are common


• Missing pinnae, closed or absent external acoustic
meatuses
• Maternal rubella causes sensorineural deafness

© 2016 Pearson Education, Inc.

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