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CHAPTER 10b
The Nervous
System:
Sensory
Systems
© 2017 Pearson Education, Ltd.
Chapter Outline

10.1 General Principles of Sensory Physiology


10.2 The Somatosensory System
10.3 Vision
10.4 The Ear and Hearing
10.5 The Ear and Equilibrium
10.6 Taste
10.7 Olfaction

© 2017 Pearson Education, Ltd.


10.3 Vision

• Anatomy of the eye


• The nature and behavior of light waves
• Accommodation
• Clinical defects in vision
• Regulating the amount of light entering the eye
• Retina
• Phototransduction
• Neural pathways for vision
• Parallel processing in the visual system
• Depth perception
© 2017 Pearson Education, Ltd.
Anatomy of the Eye

• Fluid filled
• Three layers
• Outermost
• Sclera and cornea
• Middle
• Choroid, ciliary body, iris, zonular fibers, lens
• Inner
• Retina, photoreceptors, fovea
• Optic disk (blind spot)

© 2017 Pearson Education, Ltd.


Figure 10.19a Anatomy of the eye.
Muscle

Cornea

Iris

Pupil Fovea

Optic disk
Anterior
segment
(aqueous
humor)
Lens

Zonular fibers Blood


vessels
Ciliary body Optic nerve
Sclera
Vitreous chamber
Choroid
(vitreous humor)
Retina

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Figure 10.19b Anatomy of the eye.

Fovea Blood vessels Optic disk

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Figure 10.20 The blind spot.

+ −
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The Nature and Behavior of Light Waves

• Light energy

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Figure 10.21 The electromagnetic spectrum.

1012 nm
Radio
750 nm

109 nm
TV

105 nm Micro-
waves

104 nm Infrared

750 nm Visible
350 nm
light
Ultra-
102 nm violet

1 nm X rays

10–2 nm Gamma
rays
350 nm

Cosmic
rays
© 2017 Pearson Education, Ltd.
The Nature and Behavior of Light Waves

• Reflection
• We perceive light waves reflected off objects
• Refraction: if not perpendicular, light waves bend
as they pass from one medium density to another
• Degree of refraction depends on three factors:
• Differences in densities
• Angle of light waves
• Curve of the cornea and lens, which refract light as it
enters eye

© 2017 Pearson Education, Ltd.


Figure 10.22 Refraction of light waves passing through different media.

© 2017 Pearson Education, Ltd.


Figure 10.23 Refraction of light waves passing through curved surfaces.

Focal point

Light Light

Focal length

Concave surface Convex surface

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Figure 10.24 Refraction of light waves in the eye.

Lens

Cornea

Light
waves

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Accommodation

• Increasing lens curvature enables the eye to focus


on near objects
• Mechanisms
• Under parasympathetic control
• Ciliary muscle contracts
• Decreased tension on zonular fibers
• Lens becomes rounder (refractive index increases)

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Figure 10.25a Focusing light from distant and near sources.

Nearly parallel
light rays

Flattened lens
for weak refraction

Viewing a distant object

© 2017 Pearson Education, Ltd.


Figure 10.25b Focusing light from distant and near sources.

Diverging
light rays

Round lens
for strong refraction

Viewing a near object


© 2017 Pearson Education, Ltd.
Figure 10.26a Mechanism of accommodation.

Ciliary
muscle

Zonular
fibers

Ciliary Lens
muscle (flat)
(relaxed)
Pupil
Zonular Cornea
fibers
(tight) Iris
Lens
No parasympathetic stimulation
Far vision of distant objects

© 2017 Pearson Education, Ltd.


Figure 10.26b Mechanism of accommodation.

Ciliary
muscle
Zonular
fibers

Ciliary Lens
muscle (rounded)
(contracted)
Pupil
Zonular (constricted)
fibers
(slack)
Cornea
Iris
+
Lens
Parasympathetic stimulation
Accommodation for near vision

© 2017 Pearson Education, Ltd.


Clinical Defects in Vision

• Emmetropia: normal vision


• Myopia: near-sightedness
• Hyperopia: far-sightedness
• Presbyopia: hardening of lens
• Cataract: discoloration of lens
• Glaucoma: increased volume of aqueous humor
• Astigmatism: irregularities of lens or cornea

© 2017 Pearson Education, Ltd.


Figure 10.27a Normal, near-sighted, and far-sighted vision.

Distant object Near object

Emmetropia

Focused Focused
on retina on retina
No accommodation Accommodation

© 2017 Pearson Education, Ltd.


Figure 10.27b Normal, near-sighted, and far-sighted vision.

Distant object Near object

Myopia

(Lens of eye too


Focused
strong for length Not focused on retina
of eyeball) on retina
No accommodation No accommodation

Myopia corrected
with concave lens
(which decreases
overall refractive
power) Focused
Focused on retina
on retina
No accommodation Accommodation

© 2017 Pearson Education, Ltd.


Figure 10.27c Normal, near-sighted, and far-sighted vision.

Distant object Near object

Hyperopia

(Lens of eye too


weak for length Focused Not focused
of eyeball) on retina on retina
Accommodation Accommodation

Hyperopia corrected
with convex lens
(which increases
overall refractive
power) Focused
Focused on retina
on retina
No accommodation Accommodation

© 2017 Pearson Education, Ltd.


Regulating the Amount of Light Entering
the Eye

• Size of the pupil


• Iris consists of two layers of smooth muscle
• Inner circular muscle: constrictor
• Outer radial muscle: dilator
• Hole opening in center: pupil

© 2017 Pearson Education, Ltd.


Figure 10.28a Regulation of the amount of light entering the eye.

Outer radial
muscle
Iris
Inner circular
muscle

Pupil

Anatomy of iris and pupil


© 2017 Pearson Education, Ltd.
Figure 10.28b Regulation of the amount of light entering the eye.

Parasympathetic
stimulation of
+ circular muscle

Pupillary constriction
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Figure 10.28c Regulation of the amount of light entering the eye.

Sympathetic
+ stimulation of
radial muscle

Pupillary dilation
© 2017 Pearson Education, Ltd.
Retina

• Neural tissue
• Contains photoreceptors
• Three layers
• Outer: photoreceptors
• Rods and cones
• Middle
• Bipolar, amacrine, and horizontal cells
• Inner: ganglion cells
• Macular degeneration

© 2017 Pearson Education, Ltd.


Figure 10.29 Anatomy of the retina.

Bipolar Sclera
Ganglion Photoreceptors Retinal
cells pigment Choroid
Axons of cells
optic nerve Cone Rod epithelium Retina
Amacrine Horizontal
cells cells Fovea
Macula lutea

Light

Optic nerve
Optic disk

Sclera

Choroid

Inner layer Middle layer Outer layer

© 2017 Pearson Education, Ltd.


Retina

• Cells of the retina


• Rods and cones are receptor cells
• Rods and cones communicate with bipolar cells
• Bipolar cells communicate with ganglion cells
• Axons of ganglion cells form the optic nerve
• Horizontal and amacrine cells provide lateral modulation

© 2017 Pearson Education, Ltd.


Figure 10.30 Distribution of rods and cones in the retina.

–90O 90O

–45O 45O
0O
Fovea Optic disk

Cones Rods
Number of photoreceptors

Rods
per mm2

Cones Optic disk

–90O –45O 0O 45O 90O

Fovea
Position in retina (relative to fovea)
© 2017 Pearson Education, Ltd.
Phototransduction

• Conversion of light energy to nerve signals


• Function of rods and cones
• Four photopigments
• Each has retinal and opsin
• Four different opsins
• One for the rods (black-and-white vision)
• Rhodopsin
• Three for the cones (color vision)
• L (red), M (green), and S (blue) opsins

© 2017 Pearson Education, Ltd.


Figure 10.31 Morphology of the photoreceptors.

Disks Outer
segment

Nuclei
Inner
segment

Synaptic vesicles Synaptic


terminals
Rod Cone
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Figure 10.32 Components of rods.

Fluid
inside Rhodopsin
disk

Disk
Phospho- membrane
diesterase
Retinal α
α
β γ
cGMP Breakdown
Cytosol Transducin

© 2017 Pearson Education, Ltd.


Phototransduction

• In rods in the dark:


• Levels of second messenger cGMP are high
• cGMP opens Na+ channels on the membrane of the
outer segment
• Na+ moves into cell
• Photoreceptor is depolarized, opening Ca2+ channels
• Ca2+ entry triggers neurotransmitter secretion
• Neurotransmitter acts on bipolar cells

© 2017 Pearson Education, Ltd.


Figure 10.33a Phototransduction of light. Slide 1

Retinal

cGMP levels
cGMP in the cytosol
are high

Na+ Sodium channels


are open
Sodium enters
the cell, causing
K+ a depolarization
that spreads from
the outer segment
to the terminal

Calcium channels
open in response
Ca2+ to depolarization

Calcium enters the


cell, triggering
exocytosis
Transmitter of transmitter

Bipolar cell Transmitter causes


graded potentials
in bipolar cell

State of photoreceptor in the dark


© 2017 Pearson Education, Ltd.
Figure 10.33a Phototransduction of light. Slide 2

Retinal

cGMP levels
cGMP in the cytosol
are high

Transmitter

Bipolar cell

State of photoreceptor in the dark


© 2017 Pearson Education, Ltd.
Figure 10.33a Phototransduction of light. Slide 3

Retinal

cGMP levels
cGMP in the cytosol
are high

Na+ Sodium channels


are open

Transmitter

Bipolar cell

State of photoreceptor in the dark


© 2017 Pearson Education, Ltd.
Figure 10.33a Phototransduction of light. Slide 4

Retinal

cGMP levels
cGMP in the cytosol
are high

Na+ Sodium channels


are open
Sodium enters
the cell, causing
K+ a depolarization
that spreads from
the outer segment
to the terminal

Transmitter

Bipolar cell

State of photoreceptor in the dark


© 2017 Pearson Education, Ltd.
Figure 10.33a Phototransduction of light. Slide 5

Retinal

cGMP levels
cGMP in the cytosol
are high

Na+ Sodium channels


are open
Sodium enters
the cell, causing
K+ a depolarization
that spreads from
the outer segment
to the terminal

Calcium channels
open in response
Ca2+ to depolarization

Calcium enters the


cell, triggering
exocytosis
Transmitter of transmitter

Bipolar cell

State of photoreceptor in the dark


© 2017 Pearson Education, Ltd.
Figure 10.33a Phototransduction of light. Slide 6

Retinal

cGMP levels
cGMP in the cytosol
are high

Na+ Sodium channels


are open
Sodium enters
the cell, causing
K+ a depolarization
that spreads from
the outer segment
to the terminal

Calcium channels
open in response
Ca2+ to depolarization

Calcium enters the


cell, triggering
exocytosis
Transmitter of transmitter

Bipolar cell Transmitter causes


graded potentials
in bipolar cell

State of photoreceptor in the dark


© 2017 Pearson Education, Ltd.
Phototransduction

• In rods in the light:


• Rhodopsin within membrane of disks
• Light absorbed by rhodopsin
• Retinal and opsin dissociate
• Bleached opsin activates transducin (G protein)
• Transducin activates phosphodiesterase
• Phosphodiesterase catalyzes cGMP
• Without cGMP, Na+ channels close
• Hyperpolarization of cell occurs by K+ leaving
• Hyperpolarization closes Ca2+ channels and secretion
stops

© 2017 Pearson Education, Ltd.


Figure 10.33b Phototransduction of light. Slide 1

Retinal Light is absorbed


by photopigment
Retinal and opsin dissociate
Transducin is activated

Na+ Phosphodiesterase is activated

Light cGMP levels in cytosol decrease


Sodium channels close

K+ With less sodium entering the


cell, the cell hyperpolarizes

Calcium channels
Ca2+ close

Transmitter release
is decreased
Graded potential
in bipolar cell gets
smaller

Response of photoreceptor to light


© 2017 Pearson Education, Ltd.
Figure 10.33b Phototransduction of light. Slide 2

Retinal Light is absorbed


by photopigment

Light

Response of photoreceptor to light


© 2017 Pearson Education, Ltd.
Figure 10.33b Phototransduction of light. Slide 3

Retinal Light is absorbed


by photopigment
Retinal and opsin dissociate
Transducin is activated

Na+ Phosphodiesterase is activated

Light cGMP levels in cytosol decrease


Sodium channels close

Response of photoreceptor to light


© 2017 Pearson Education, Ltd.
Figure 10.33b Phototransduction of light. Slide 4

Retinal Light is absorbed


by photopigment
Retinal and opsin dissociate
Transducin is activated

Na+ Phosphodiesterase is activated

Light cGMP levels in cytosol decrease


Sodium channels close

K+ With less sodium entering the


cell, the cell hyperpolarizes

Response of photoreceptor to light


© 2017 Pearson Education, Ltd.
Figure 10.33b Phototransduction of light. Slide 5

Retinal Light is absorbed


by photopigment
Retinal and opsin dissociate
Transducin is activated

Na+ Phosphodiesterase is activated

Light cGMP levels in cytosol decrease


Sodium channels close

K+ With less sodium entering the


cell, the cell hyperpolarizes

Calcium channels
Ca2+ close

Response of photoreceptor to light


© 2017 Pearson Education, Ltd.
Figure 10.33b Phototransduction of light. Slide 6

Retinal Light is absorbed


by photopigment
Retinal and opsin dissociate
Transducin is activated

Na+ Phosphodiesterase is activated

Light cGMP levels in cytosol decrease


Sodium channels close

K+ With less sodium entering the


cell, the cell hyperpolarizes

Calcium channels
Ca2+ close

Transmitter release
is decreased
Graded potential
in bipolar cell gets
smaller

Response of photoreceptor to light


© 2017 Pearson Education, Ltd.
Rod and Cone Sensitivity

• Scotopic vision: rods


• Mesopic vision: rods and cones
• Photopic vision: cones

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Figure 10.34 Absorbance spectra for the different photoreceptors.
L
Rods cones
S M
cones cones

100
Light absorbance
(% of maximum)

75

50

25

400 430 500 530 560 600


Wavelength (nm)
© 2017 Pearson Education, Ltd.
Figure 10.35 Rod and cone sensitivity.

–6 –4 –2 0 2 4 6 8
Luminance
(log cd/cm2)

Indoor
Starlight Moonlight lighting Sunlight
Luminance of
natural light
on white

Scotopic Mesopic Photopic


Type of vision

Highest
visual acuity

© 2017 Pearson Education, Ltd.


© 2017 Pearson Education, Ltd.
Color Vision

• Three types of cones:


• Blue: responds best to 420 nm
• Green: responds best to 530 nm
• Red: responds best to 560 nm
• Each type responds best (not exclusively) to its
color

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Color Vision

• Color perception
• Brain compares responses of cones
• Cone comparison = color perception
• Opponent-process theory
• Some colors inhibit perception of others
• Red–green/green–red
• Blue–yellow/yellow–blue
• Black–white/white–black

© 2017 Pearson Education, Ltd.


Figure 10.36 Theoretical state if only two types of cones existed.

P1 P2
Absorbance

450 500 550


Wavelength (nm)
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Color Vision

• Color adaptation experiment


• Stare at the next slide
• Continue staring at the same place on the screen
with the following slide

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Figure 10.37 Opponent-color adaptation.

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Color Vision

• Color blindness
• Red-green most common
• Affects 4% of men
• Ishihara charts

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Clinical Connections 10.2

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Light Input to Circadian Rhythms

• Many daily rhythms


• Without light, circadian rhythms would run longer
than 24 hours
• Special photoreceptors contain melanopsin
• Link to rhythm-generating center of the brain

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Bleaching of Photoreceptors in Light

• Adaptation to light and dark


• Small changes in light intensity
• Pupillary dilation and constriction
• Larger changes in light intensity
• Changes in photopigment
• Bleaching

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Bleaching of Photoreceptors in Light

• Going from light to dark


• Exposure to light "bleaches" the rods
• Opsin separated from retinal
• No more light can be absorbed
• Move to dark
• Sensitivity of rods is low due to previous "bleaching"
• Retinal and opsin reassociate
• Can again detect light

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Bleaching of Photoreceptors in Light

• Going from dark to light


• Exposure to dark maintains rods in most sensitive state
• Opsin and retinal are associated
• Readily absorb light
• Move to bright light
• Sensitive rods are overwhelmed
• Rods begin to be "bleached"
• Can again see clearly

© 2017 Pearson Education, Ltd.


Neural Processing in the Retina

• Photoreceptors communicate to bipolar cells, and


bipolar cells communicate to ganglion cells
• Convergence
• More than one photoreceptor to bipolar neuron
• More than one bipolar cell to ganglion cell

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Neural Processing in the Retina

• Rods converge more


• Lower visual acuity
• Greater sensitivity
• In fovea
• One cone communicates with one bipolar cell
• Greater acuity
• Lower sensitivity

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Neural Processing in the Retina

• Bipolar cell receptive fields


• Glutamate
• Stimulatory at ionotropic receptors
• Inhibitory at metabotropic receptors
• Direct photoreceptor → bipolar causes response in
center receptive field
• Glutamate released in dark (decreased in light)
• Bipolars excited by glutamate are inhibited by light
• OFF bipolar cell
• Bipolars inhibited by glutamate are excited by light
• ON bipolar cell

© 2017 Pearson Education, Ltd.


Neural Processing in the Retina

• Bipolar cell receptive fields


• OFF bipolar cell
• Excited by light in surround receptive field
• Due to interaction between photoreceptors, bipolar cells
and horizontal cells
• ON bipolar cell
• Inhibited by light in surround receptive field
• More correct terminology: ON-center, OFF-surround
and OFF-center, ON-surround

© 2017 Pearson Education, Ltd.


Figure 10.38 Receptive fields of ganglion cells.

+
+ +
+
+ +
+
+ + +
+
+ + + +
+ + + +
+
+ +
+

ON-center, OFF-surround OFF-center, ON-surround


ganglion cell ganglion cell

© 2017 Pearson Education, Ltd.


Neural Processing in the Retina

• Ganglion cell receptive field


• Bipolar cells transmit graded potentials, not action
potentials
• Receptive field properties of bipolar cells are continuous
to ganglion cells
• ON-center, OFF-surround and OFF-center, ON-surround
• Lateral inhibition by amacrine cells
• Transmit action potentials
• Ganglion cell axons make up the optic nerve
• Disinhibition causes excitation

© 2017 Pearson Education, Ltd.


Figure 10.39 Neural circuitry resulting in an ON-center, OFF-surround ganglion cell.

Surround
Center

Surround Center Surround


cone cone cone

– –

Bipolar
cell

ON-center
OFF-surround To optic nerve
ganglion cell
© 2017 Pearson Education, Ltd.
Neural Pathways for Vision

• Ganglionic cells
• Optic nerve (cranial nerve II)
• Optic chiasm
• Optic tract
• Lateral geniculate body of thalamus synapses
• Optic radiations
• Visual cortex synapses
• Right visual field to left cortex, and vice versa

© 2017 Pearson Education, Ltd.


Figure 10.40 Neural pathways for vision.
Binocular visual field

Nasal retina
Temporal retina

Optic nerve

Optic chiasm

Optic tract

Lateral geniculate
body of thalamus

Optic radiations

Primary visual
cortex
© 2017 Pearson Education, Ltd.
Parallel Processing in the Visual System

• Parallel pathways transfer different types of visual


information
• Color
• Shape
• Movement

© 2017 Pearson Education, Ltd.


Depth Perception

• Binocular visual field


• Two perspectives of visual field, one from each
eye
• Brain constructs three-dimensional image

© 2017 Pearson Education, Ltd.

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