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Chapter 49

Sensory and Motor


Mechanisms

PowerPoint Lectures for


Biology, Seventh Edition
Neil Campbell and Jane Reece

Lectures by Chris Romero


Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Overview: Sensing and Acting

• Bats use sonar to detect their prey

• Moths, a common prey for bats


– Can detect the bat’s sonar and attempt to flee

Figure 49.1
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• Both of these organisms
– Have complex sensory systems that facilitate
their survival

• The structures that make up these systems


– Have been transformed by evolution into
diverse mechanisms that sense various stimuli
and generate the appropriate physical
movement

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• Concept 49.1: Sensory receptors transduce
stimulus energy and transmit signals to the
central nervous system
• Sensations are action potentials
– That reach the brain via sensory neurons

• Once the brain is aware of sensations


– It interprets them, giving the perception of
stimuli

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• Sensations and perceptions
– Begin with sensory reception, the detection of
stimuli by sensory receptors

• Exteroreceptors
– Detect stimuli coming from the outside of the
body

• Interoreceptors
– Detect internal stimuli

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Functions Performed by Sensory Receptors
• All stimuli represent forms of energy

• Sensation involves converting this energy


– Into a change in the membrane potential of
sensory receptors

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• Sensory receptors perform four functions in this
process
– Sensory transduction, amplification,
transmission, and integration

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• Two types of sensory receptors exhibit these
functions
– A stretch receptor in a crayfish

Weak Strong
Muscle
muscle stretch muscle stretch
Dendrites
–50 Receptor potential –50
potential (mV)

–70 –70
Membrane

Stretch
receptor Action potentials
0 0
Axon
–70 –70
0 1 2 3 4 5 6 7 01 2 3 4 5 67
Time (sec) Time (sec)
(a) Crayfish stretch receptors have dendrites stretch, producing a receptor potential in the receptor. A stronger stretch produces
embedded in abdominal muscles. When the stretch receptor. The receptor potential triggers a larger receptor potential and higher
abdomen bends, muscles and dendrites action potentials in the axon of the stretch requency of action potentials.

Figure 49.2a

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– A hair cell found in vertebrates

“Hairs” of No fluid Fluid moving in Fluid moving in


hair cell movement one direction other direction
Neuro- More
trans- Less
neuro-
mitter at neuro-
trans-
synapse trans-
mitter
mitter
Axon –50 –50 Receptor potential –50

potential (mV)

potential (mV)
potential (mV)

Membrane
–70

Membrane
–70 –70
Membrane

Action potentials
0 0 0

–70 –70 –70


01 2 3 4 5 6 7 0 1 2 3 4 5 6 7 01 2 3 4 5 6 7
Time (sec) Time (sec) Time (sec)

(b) Vertebrate hair cells have specialized cilia with a sensory neuron, which conducts action of action potentials in the sensory neuron.
or microvilli (“hairs”) that bend when sur- potentials to the CNS. Bending in one direction Bending in the other direction has the opposite
rounding fluid moves. Each hair cell releases depolarizes the hair cell, causing it to release effects. Thus, hair cells respond to the direction
an excitatory neurotransmitter at a synapse more neurotransmitter and increasing frequency of motion as well as to its strength and speed.s

Figure 49.2b

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Sensory Transduction
• Sensory transduction is the conversion of
stimulus energy
– Into a change in the membrane potential of a
sensory receptor

• This change in the membrane potential


– Is known as a receptor potential

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• Many sensory receptors are extremely
sensitive
– With the ability to detect the smallest physical
unit of stimulus possible

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Amplification
• Amplification is the strengthening of stimulus
energy
– By cells in sensory pathways

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Transmission
• After energy in a stimulus has been transduced
into a receptor potential
– Some sensory cells generate action potentials,
which are transmitted to the CNS

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• Sensory cells without axons
– Release neurotransmitters at synapses with
sensory neurons

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Integration
• The integration of sensory information
– Begins as soon as the information is received

– Occurs at all levels of the nervous system

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• Some receptor potentials
– Are integrated through summation

• Another type of integration is sensory


adaptation
– A decrease in responsiveness during
continued stimulation

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Types of Sensory Receptors
• Based on the energy they transduce, sensory
receptors fall into five categories
– Mechanoreceptors

– Chemoreceptors

– Electromagnetic receptors

– Thermoreceptors

– Pain receptors

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Mechanoreceptors
• Mechanoreceptors sense physical deformation
– Caused by stimuli such as pressure, stretch,
motion, and sound

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• The mammalian sense of touch
– Relies on mechanoreceptors that are the
dendrites of sensory neurons
Cold
Light touch Pain Hair
Heat

Epidermis

Dermis

Figure 49.3 Nerve Connective tissue Hair movement Strong pressure

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Chemoreceptors
• Chemoreceptors include
– General receptors that transmit information
about the total solute concentration of a
solution
– Specific receptors that respond to individual
kinds of molecules

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• Two of the most sensitive and specific
chemoreceptors known
– Are present in the antennae of the male
silkworm moth

Figure 49.4 0.1 mm


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Electromagnetic Receptors
• Electromagnetic receptors detect various forms
of electromagnetic energy
– Such as visible light, electricity, and
magnetism

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• Some snakes have very sensitive infrared
receptors
– That detect body heat of prey against a colder
background

Figure 49.5a
(a) This rattlesnake and other pit vipers have a pair of infrared receptors,
one between each eye and nostril. The organs are sensitive enough
to detect the infrared radiation emitted by a warm mouse a meter away.
The snake moves its head from side to side until the radiation is detected
equally by the two receptors, indicating that the mouse is straight ahead.

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• Many mammals appear to use the Earth’s
magnetic field lines
– To orient themselves as they migrate

Figure 49.5b
(b) Some migrating animals, such as these beluga whales, apparently
sense Earth’s magnetic field and use the information, along with
other cues, for orientation.

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Thermoreceptors
• Thermoreceptors, which respond to heat or
cold
– Help regulate body temperature by signaling
both surface and body core temperature

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Pain Receptors
• In humans, pain receptors, also called
nociceptors
– Are a class of naked dendrites in the epidermis

– Respond to excess heat, pressure, or specific


classes of chemicals released from damaged
or inflamed tissues

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• Concept 49.2: The mechanoreceptors involved
with hearing and equilibrium detect settling
particles or moving fluid
• Hearing and the perception of body equilibrium
– Are related in most animals

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Sensing Gravity and Sound in Invertebrates
• Most invertebrates have sensory organs called
statocysts
– That contain mechanoreceptors and function in
their sense of equilibrium

Ciliated
receptor cells

Statolith Cilia

Figure 49.6 Sensory nerve fibers

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• Many arthropods sense sounds with body hairs
that vibrate
– Or with localized “ears” consisting of a
tympanic membrane and receptor cells

Tympanic
membrane

Figure 49.7

1 mm
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Hearing and Equilibrium in Mammals
• In most terrestrial vertebrates
– The sensory organs for hearing and
equilibrium are closely associated in the ear

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• Exploring the structure of the human ear
1 Overview of ear structure 2 The middle ear and inner ear

Incus
Skull Semicircular
bones canals
Stapes
Middle
Outer ear ear Inner ear Malleus
Auditory nerve,
to brain

Pinna

Tympanic
membrane

Cochlea
Eustachian
Auditory tube
canal

Oval Eustachian
Tympanic window tube
Hair cells Tectorial
membrane
membrane Round
window

Cochlear duct
Bone

Vestibular canal

Auditory nerve
Basilar Axons of To auditory
membrane sensory neurons nerve Tympanic canal

Organ of Corti
Figure 49.8 4 The organ of Corti 3 The cochlea

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Hearing
• Vibrating objects create percussion waves in
the air
– That cause the tympanic membrane to vibrate

• The three bones of the middle ear


– Transmit the vibrations to the oval window on
the cochlea

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• These vibrations create pressure waves in the
fluid in the cochlea
– That travel through the vestibular canal and
ultimately strike the round window

Cochlea

Stapes Axons of
sensory
neurons
Oval
window Vestibular
canal Perilymph Apex

Base

Round Tympanic
window canal Basilar
Figure 49.9 membrane

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• The pressure waves in the vestibular canal
– Cause the basilar membrane to vibrate up and
down causing its hair cells to bend

• The bending of the hair cells depolarizes their


membranes
– Sending action potentials that travel via the
auditory nerve to the brain

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• The cochlea can distinguish pitch
– Because the basilar membrane is not uniform
along its length
Cochlea
(uncoiled)
Apex
(wide and flexible)
Basilar
membrane

500 Hz
1 kHz (low pitch)

2 kHz

4 kHz

8 kHz

16 kHz Frequency producing maximum


(high pitch) vibration
Base
Figure 49.10 (narrow and stiff)

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• Each region of the basilar membrane vibrates
most vigorously
– At a particular frequency and leads to
excitation of a specific auditory area of the
cerebral cortex

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Equilibrium
• Several of the organs of the inner ear
– Detect body position and balance

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• The utricle, saccule, and semicircular canals in
the inner ear
– Function in balance and equilibrium
The semicircular canals, arranged in three Each canal has at its base a When the head changes its rate
spatial planes, detect angular movements swelling called an ampulla, of rotation, inertia prevents
of the head. containing a cluster of hair cells. endolymph in the semicircular
canals from moving with the head,
so the endolymph presses against
the cupula, bending the hairs.
Flow
of endolymph
Flow
of endolymph
Vestibular nerve

Cupula
Hairs

Hair
cell

Vestibule Nerve
fibers
Utricle Body movement
Saccule

Figure 49.11 The utricle and saccule tell the brain which The hairs of the hair cells Bending of the hairs increases the
way is up and inform it of the body’s project into a gelatinous cap frequency of action potentials in
position or linear acceleration. called the cupula. sensory neurons in direct
proportion to the amount of
rotational acceleration.

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Hearing and Equilibrium in Other Vertebrates
• Like other vertebrates, fishes and amphibians
– Also have inner ears located near the brain

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• Most fishes and aquatic amphibians
– Also have a lateral line system along both
sides of their body

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• The lateral line system contains
mechanoreceptors
– With hair cells that respond to water movement
Lateral
line

Opening of lateral
Lateral line canal
Scale line canal
Epidermis Neuromast

Segmental muscles of body wall Lateral nerve

Cupula

Sensory
hairs
Supporting cell Hair cell

Figure 49.12 Nerve fiber

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• Concept 49.3: The senses of taste and smell
are closely related in most animals
• The perceptions of gustation (taste) and
olfaction (smell)
– Are both dependent on chemoreceptors that
detect specific chemicals in the environment

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• The taste receptors of insects are located
within sensory hairs called sensilla
– Which are located on the feet and in
mouthparts

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EXPERIMENT Insects taste using gustatory sensilla (hairs) on their feet and To brain
mouthparts. Each sensillum contains four chemoreceptors with dendrites that
extend to a pore at the tip of the sensillum. To study the sensitivity of each
chemoreceptor, researchers immobilized a blowfly (Phormia regina) by attaching Chemo-
receptors Sensillum
it to a rod with wax. They then inserted the tip of a microelectrode into one
sensillum to record action potentials in the chemoreceptors, while they used a
pipette to touch the pore with various test substances. Microelectrode

To voltage
recorder

RESULTS Each chemoreceptor is especially sensitive to a particular


class of substance, but this specificity is relative; each cell can respond to Pore at tip
some extent to a broad range of different chemical stimuli.
Pipette containing
test substance

Number of action potentials


Chemoreceptors

in first second of response


50

30
CONCLUSION Any natural food probably stimulates multiple chemoreceptors. By
integrating sensations, the insect’s brain can apparently distinguish a very large
number of tastes. 10
0
0.5 M Meat 0.5 M Honey
NaCl Sucrose
Figure 49.13 Stimulus

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Taste in Humans
• The receptor cells for taste in humans
– Are modified epithelial cells organized into
taste buds

• Five taste perceptions involve several signal


transduction mechanisms
– Sweet, sour, salty, bitter, and umami (elicited
by glutamate)

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• Transduction in taste receptors
– Occurs by several mechanisms
Taste pore Sugar molecule

Taste bud
Sensory
receptor
cells

Tongue Sensory
neuron

1 A sugar molecule binds


to a receptor protein on
Sugar the sensory receptor cell.

Sugar G protein Adenylyl cyclase


receptor
2 Binding initiates a signal transduction pathway
involving cyclic AMP and protein kinase A.

ATP
cAMP

Protein
kinase A 3 Activated protein kinase A closes K+ channels in
the membrane.
SENSORY
K+
RECEPTOR 4 The decrease in the membrane’s permeability to
CELL Synaptic K+ depolarizes the membrane.
vesicle

—Ca2+

5 Depolarization opens voltage-gated calcium ion


(Ca2+) channels, and Ca2+ diffuses into the receptor
Neurotransmitter cell.

6 The increased Ca2+ concentration causes


synaptic vesicles to release neurotransmitter.

Figure 49.14 Sensory neuron

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Smell in Humans
• Olfactory receptor cells
– Are neurons that line the upper portion of the
nasal cavity

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• When odorant molecules bind to specific
receptors
– A signal transduction pathway is triggered,
sending action potentials to the brain
Brain
Action potentials

Odorant Olfactory bulb

Nasal cavity
Bone

Epithelial cell

Odorant
receptors Chemoreceptor

Plasma
membrane Cilia

Figure 49.15 Odorant Mucus

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• Concept 49.4: Similar mechanisms underlie
vision throughout the animal kingdom
• Many types of light detectors
– Have evolved in the animal kingdom and may
be homologous

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Vision in Invertebrates
• Most invertebrates
– Have some sort of light-detecting organ

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• One of the simplest is the eye cup of
planarians
– Which provides information about light
intensity and direction but does not form
images Light

Light shining from


the front is detected

Photoreceptor
Nerve to
Visual pigment brain
Screening
Ocellus pigment

Light shining from


behind is blocked
Figure 49.16 by the screening pigment

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• Two major types of image-forming eyes have
evolved in invertebrates
– The compound eye and the single-lens eye

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• Compound eyes are found in insects and
crustaceans
– And consist of up to several thousand light
detectors called ommatidia
(a) The faceted eyes on the
head of a fly,
photographed with
a stereomicroscope.

2 mm
(b) The cornea and crystalline cone of
each ommatidium function as Cornea
a lens that focuses light on the
rhabdom, a stack of pigmented
plates inside a circle of Crystalline Lens
photoreceptors. The rhabdom cone
traps light and guides it to
photoreceptors. The image
formed by a compound eye is a
mosaic of dots produced by different Rhabdom
intensities of light entering the
many ommatidia from different angles. Photoreceptor
Axons
Figure 49.17a–b
Ommatidium
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• Single-lens eyes
– Are found in some jellies, polychaetes, spiders,
and many molluscs
– Work on a camera-like principle

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The Vertebrate Visual System
• The eyes of vertebrates are camera-like
– But they evolved independently and differ from
the single-lens eyes of invertebrates

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Structure of the Eye
• The main parts of the vertebrate eye are
– The sclera, which includes the cornea

– The choroid, a pigmented layer

– The conjunctiva, that covers the outer surface


of the sclera

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– The iris, which regulates the pupil

– The retina, which contains photoreceptors

– The lens, which focuses light on the retina

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• The structure of the vertebrate eye
Sclera Choroid

Retina
Ciliary body

Fovea (center
Suspensory of visual field)
ligament

Cornea

Iris
Optic
Pupil nerve

Aqueous
humor

Lens

Vitreous humor
Central artery and
vein of the retina

Optic disk
Figure 49.18 (blind spot)
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• Humans and other mammals
– Focus light by changing the shape of the lens
Front view of lens
Ciliary muscles contract, pulling and ciliary muscle
border of choroid toward lens Lens (rounder)
Choroid

Suspensory ligaments relax Retina

Ciliary
muscle

Lens becomes thicker and rounder, Suspensory


focusing on near objects ligaments

(a) Near vision (accommodation)

Ciliary muscles relax, and border of


choroid moves away from lens Lens (flatter)

Suspensory ligaments
pull against lens

Lens becomes flatter, focusing on


distant objects

Figure 49.19a–b (b) Distance vision

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• The human retina contains two types of
photoreceptors
– Rods are sensitive to light but do not
distinguish colors
– Cones distinguish colors but are not as
sensitive

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Sensory Transduction in the Eye
• Each rod or cone in the vertebrate retina
– Contains visual pigments that consist of a light-
absorbing molecule called retinal bonded to a
protein called opsin

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• Rods contain the pigment rhodopsin
– Which changes shape when it absorbs light
Rod

Outer
segment
H
H C O
H CH3
C CH3 H3C C H
H2C C H H C
H2C C C C C
Disks
C C C C H
CH3 H CH3 H
cis isomer
Cell body Inside
of disk
Light Enzymes

Synaptic
terminal H
H CH3
C CH3
H2C C H H H H
H2C C C C C C O
C C C C C C
H
Cytosol CH3 H CH3 CH3 CH3

Retinal trans isomer


Rhodopsin
Opsin
(b) Retinal exists as two isomers. Absorption of light converts
(a) Rods contain the visual pigment rhodopsin, which is embedded in the cis isomer to the trans isomer, which
a stack of membranous disks in the rod’s outer segment. causes opsin to change its conformation (shape).
Rhodopsin consists of the light-absorbing molecule retinal After a few minutes, retinal detaches from opsin.
bonded to opsin, a protein. Opsin has seven α helices that span In the dark, enzymes convert retinal back to its cis
form, which recombines with opsin to form rhodopsin.
Figure 49.20a, b the disk membrane.

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Processing Visual Information
• The processing of visual information
– Begins in the retina itself

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• Absorption of light by retinal
– Triggers a signal transduction pathway

Light EXTRACELLULAR
INSIDE OF DISK
FLUID
Active rhodopsin PDE

Membrane
potential (mV)
Plasma
CYTOSOL
membrane 0
Dark Light
cGMP
Inactive rhodopsin Transducin Disk membrane
– 40
GMP
Na+ – Hyper-
– 70 polarization
2 Active
1 Light 3 Transducin 4 Activated PDE
rhodopsin Time
isomerizes activates the detaches cyclic
in turn
retinal, which enzyme guanosine
activates a G 5 The Na+ channels
activates phos- monophosphate
protein close when cGMP
rhodopsin. phodiesterae (cGMP) from
called detaches. The
(PDE). Na+ channels in
transducin. membrane’s
the plasma
membrane by permeability to
Na+ Na+ decreases,
hydrolyzing
cGMP to GMP. and the rod
hyperpolarizes.
Figure 49.21

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• In the dark, both rods and cones
– Release the neurotransmitter glutamate into
the synapses with neurons called bipolar cells,
which are either hyperpolarized or depolarized

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• In the light, rods and cones hyperpolarize
– Shutting off their release of glutamate

• The bipolar cells


– Are then either depolarized or hyperpolarized
Dark Responses Light Responses

Rhodopsin inactive Rhodopsin active

Na+ channels open Na+ channels closed

Rod depolarized Rod hyperpolarized

Glutamate No glutamate
released released

Bipolar cell either Bipolar cell either


depolarized or hyperpolarized or
hyperpolarized, depolarized,
depending on depending on
glutamate receptors glutamate receptors
Figure 49.22
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• Three other types of neurons contribute to
information processing in the retina
– Ganglion cells, horizontal cells, and amacrine
cells
Retina
Optic nerve

To
brain

Retina

Photoreceptors
Neurons
Cone Rod

Amacrine
cell Horizontal
Optic cell
nerve Pigmented
Figure 49.23 fibers
Ganglion
cell
Bipolar
cell epithelium

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• Signals from rods and cones
– Travel from bipolar cells to ganglion cells

• The axons of ganglion cells are part of the optic


nerve
– That transmit information to the brain
Left Right
visual visual
field field

Left Right
eye eye
Optic nerve

Optic chiasm

Lateral
geniculate
nucleus

Primary
Figure 49.24 visual cortex

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• Most ganglion cell axons lead to the lateral
geniculate nuclei of the thalamus
– Which relays information to the primary visual
cortex

• Several integrating centers in the cerebral


cortex
– Are active in creating visual perceptions

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• Concept 49.5: Animal skeletons function in
support, protection, and movement
• The various types of animal movements
– All result from muscles working against some
type of skeleton

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Types of Skeletons
• The three main functions of a skeleton are
– Support, protection, and movement

• The three main types of skeletons are


– Hydrostatic skeletons, exoskeletons, and
endoskeletons

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Hydrostatic Skeletons
• A hydrostatic skeleton
– Consists of fluid held under pressure in a
closed body compartment

• This is the main type of skeleton


– In most cnidarians, flatworms, nematodes, and
annelids

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• Annelids use their hydrostatic skeleton for
peristalsis
– A type of movement on land produced by
rhythmic waves of muscle contractions
Longitudinal Circular Circular Longitudinal
(a) Body segments at the head and just in front muscle relaxed muscle muscle muscle
of the rear are short and thick (longitudinal (extended) contracted relaxed contracted
muscles contracted; circular muscles
relaxed) and anchored to the ground by
bristles. The other segments are thin and
elongated (circular muscles contracted;
longitudinal muscles relaxed.)

Bristles Head

(b) The head has moved forward because


circular muscles in the head segments have
contracted. Segments behind the head and
at the rear are now thick and anchored, thus
preventing the worm from slipping backward.

(c) The head segments are thick again and


anchored in their new positions. The rear
segments have released their hold on the
Figure 49.25a–c ground and have been pulled forward.

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Exoskeletons
• An exoskeleton is a hard encasement
– Deposited on the surface of an animal

• Exoskeletons
– Are found in most molluscs and arthropods

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Endoskeletons
• An endoskeleton consists of hard supporting
elements
– Such as bones, buried within the soft tissue of
an animal

• Endoskeletons
– Are found in sponges, echinoderms, and
chordates

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• The mammalian skeleton is built from more
than 200 bones
– Some fused together and others connected at
joints by ligaments that allow freedom of
movement

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• The human skeleton
key Head of
Axial skeleton
Examples humerus
Skull of joints
Appendicular
Scapula
skeleton
1
Shoulder Clavicle
girdle Scapula

Sternum
Rib 1 Ball-and-socket joints, where the humerus contacts
2 the shoulder girdle and where the femur contacts the
Humerus

Vertebra 3 pelvic girdle, enable us to rotate our arms and


Radius legs and move them in several planes.
Ulna Humerus
Pelvic
girdle
Carpals

Phalanges
Ulna
Metacarpals
Femur 2 Hinge joints, such as between the humerus and
Patella the head of the ulna, restrict movement to a single
plane.
Tibia

Fibula

Ulna
Tarsals Radius
Figure 49.26 Metatarsals
Phalanges 3 Pivot joints allow us to rotate our forearm at the
elbow and to move our head from side to side.
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Physical Support on Land
• In addition to the skeleton
– Muscles and tendons help support large land
vertebrates

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• Concept 49.6: Muscles move skeletal parts by
contracting
• The action of a muscle
– Is always to contract

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• Skeletal muscles are attached to the skeleton
in antagonistic pairs
– With each member of the pair working against
each other Human Grasshopper
Extensor
muscle
Biceps relaxes
Tibia
contracts
flexes

Flexor
muscle
Triceps Forearm contracts
relaxes flexes

Extensor
Biceps muscle Tibia
relaxes contracts extends

Forearm
extends Flexor
muscle
Triceps relaxes
Figure 49.27 contracts

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Vertebrate Skeletal Muscle
• Vertebrate skeletal muscle
– Is characterized by a hierarchy of smaller and
Muscle

smaller units
Bundle of
muscle fibers
Nuclei

Single muscle fiber


(cell)
Plasma membrane
Myofibril Z line

Light
band Dark band

Sarcomere

TEM 0.5 µm
I band A band I band
M line
Thick
filaments
(myosin)
Thin
filaments
Figure 49.28 (actin)

Z line H zone Z line


Sarcomere

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• A skeletal muscle consists of a bundle of long
fibers
– Running parallel to the length of the muscle

• A muscle fiber
– Is itself a bundle of smaller myofibrils arranged
longitudinally

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• The myofibrils are composed to two kinds of
myofilaments
– Thin filaments, consisting of two strands of
actin and one strand of regulatory protein
– Thick filaments, staggered arrays of myosin
molecules

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• Skeletal muscle is also called striated muscle
– Because the regular arrangement of the
myofilaments creates a pattern of light and
dark bands

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• Each repeating unit is a sarcomere
– Bordered by Z lines

• The areas that contain the myofilments


– Are the I band, A band, and H zone

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The Sliding-Filament Model of Muscle Contraction
• According to the sliding-filament model of
muscle contraction
– The filaments slide past each other
longitudinally, producing more overlap
between the thin and thick filaments

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• As a result of this sliding
– The I band and the H zone shrink
0.5 µm

(a) Relaxed muscle fiber. In a relaxed muscle fiber, the I bands Z H


and H zone are relatively wide. A
Sarcomere

(b) Contracting muscle fiber. During contraction, the thick and


thin filaments slide past each other, reducing the width of the
I bands and H zone and shortening the sarcomere.

(c) Fully contracted muscle fiber. In a fully contracted muscle


fiber, the sarcomere is shorter still. The thin filaments overlap,
eliminating the H zone. The I bands disappear as the ends of
Figure 49.29a–c the thick filaments contact the Z lines.

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• The sliding of filaments is based on
– The interaction between the actin and myosin
molecules of the thick and thin filaments

• The “head” of a myosin molecule binds to an


actin filament
– Forming a cross-bridge and pulling the thin
filament toward the center of the sarcomere

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• Myosin-actin interactions underlying muscle
fiber contraction
Thick filament
1 Starting here, the myosin head is
Thin filaments bound to ATP and is in its low-
energy confinguration.

5 Binding of a new mole-


cule of ATP releases the Thin filament
myosin head from actin, Myosin head (low-
and a new cycle begins. ATP
energy configuration) 2 The myosin head hydrolyzes
ATP ATP to ADP and inorganic
phosphate ( P I ) and is in its
high-energy configuration.
Cross-bridge
Thin filament moves Thick binding site
Actin
toward center of sarcomere. filament

Myosin head (low- ADP Myosin head (high-


Pi
energy configuration) energy configuration)

13 The myosin head binds to


ADP actin, forming a cross-
ADP + Pi Pi Cross-bridge bridge.
4 Releasing ADP and ( P i), myosin
relaxes to its low-energy configuration,
Figure 49.30 sliding the thin filament.
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The Role of Calcium and Regulatory Proteins
• A skeletal muscle fiber contracts
– Only when stimulated by a motor neuron

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• When a muscle is at rest
– The myosin-binding sites on the thin filament
are blocked by the regulatory protein
tropomyosin

Tropomyosin Ca2+-binding sites


Actin
Troponin complex

Figure 49.31a (a) Myosin-binding sites blocked

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• For a muscle fiber to contract
– The myosin-binding sites must be uncovered

• This occurs when calcium ions (Ca2+)


– Bind to another set of regulatory proteins, the
troponin complex
Ca2+

Myosin-
binding site

Figure 49.31b (b) Myosin-binding sites exposed

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• The stimulus leading to the contraction of a
skeletal muscle fiber
– Is an action potential in a motor neuron that
makes a synapse with the muscle fiber
Motor Mitochondrion
neuron axon

Synaptic
terminal

T tubule

Sarcoplasmic
reticulum Ca2+ released
from sarcoplasmic
reticulum
Myofibril
Sarcomere
Plasma membrane
Figure 49.32 of muscle fiber

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• The synaptic terminal of the motor neuron
– Releases the neurotransmitter acetylcholine,
depolarizing the muscle and causing it to
produce an action potential

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• Action potentials travel to the interior of the
muscle fiber
– Along infoldings of the plasma membrane
called transverse (T) tubules
• The action potential along the T tubules
– Causes the sarcoplasmic reticulum to release
Ca2+
• The Ca2+ binds to the troponin-tropomyosin
complex on the thin filaments
– Exposing the myosin-binding sites and
allowing the cross-bridge cycle to proceed
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Review of contraction in a skeletal muscle fiber
Synaptic 1 Acetylcholine (ACh) released by synaptic terminal diffuses across synaptic
terminal cleft and binds to receptor proteins on muscle fiber’s plasma membrane,
of motor triggering an action potential in muscle fiber.
neuron
PLASMA MEMBRANE
Synaptic cleft T TUBULE

2 Action potential is propa-


ACh gated along plasma
membrane and down SR
T tubules.

3 Action potential
triggers Ca2+
release from sarco-
Ca2+ plasmic reticulum
(SR).

7 Tropomyosin blockage of myosin-


binding sites is restored; contraction
4 Calcium ions bind to troponin;
ends, and muscle fiber relaxes. troponin changes shape,
removing blocking action
Ca2+ of tropomyosin; myosin-binding
sites exposed.
CYTOSOL

6 Cytosolic Ca is
2+

removed by active
ADP transport into
P2 SR after action
potential ends.

5 Myosin cross-bridges alternately attach


to actin and detach, pulling actin
filaments toward center of sarcomere;
Figure 49.33 ATP powers sliding of filaments.

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Neural Control of Muscle Tension
• Contraction of a whole muscle is graded
– Which means that we can voluntarily alter the
extent and strength of its contraction

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• There are two basic mechanisms by which the
nervous system produces graded contractions
of whole muscles
– By varying the number of fibers that contract

– By varying the rate at which muscle fibers are


stimulated

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• In a vertebrate skeletal muscle
– Each branched muscle fiber is innervated by
only one motor neuron

• Each motor neuron


– May synapse with multiple muscle fibers
Motor Motor
unit 1 unit 2
Spinal cord Synaptic terminals

Nerve

Motor neuron
cell body

Motor neuron
axon

Muscle

Muscle fibers

Figure 49.34 Tendon

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• A motor unit
– Consists of a single motor neuron and all the
muscle fibers it controls

• Recruitment of multiple motor neurons


– Results in stronger contractions

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• A twitch
– Results from a single action potential in a
motor neuron

• More rapidly delivered action potentials


– Produce a graded contraction by summation
Tetanus
Tension

Summation of
two twitches
Single
twitch

Time
Action
potential Pair of
Series of action
action
potentials at
potentials
high frequency
Figure 49.35
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• Tetanus is a state of smooth and sustained
contraction
– Produced when motor neurons deliver a volley
of action potentials

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Types of Muscle Fibers
• Skeletal muscle fibers are classified as slow
oxidative, fast oxidative, and fast glycolytic
– Based on their contraction speed and major
pathway for producing ATP

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• Types of skeletal muscles

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Other Types of Muscle
• Cardiac muscle, found only in the heart
– Consists of striated cells that are electrically
connected by intercalated discs
– Can generate action potentials without neural
input

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• In smooth muscle, found mainly in the walls of
hollow organs
– The contractions are relatively slow and may
be initiated by the muscles themselves

• In addition, contractions may be caused by


– Stimulation from neurons in the autonomic
nervous system

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• Concept 49.7: Locomotion requires energy to
overcome friction and gravity
• Movement is a hallmark of all animals
– And usually necessary for finding food or
evading predators

• Locomotion
– Is active travel from place to place

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Swimming
• Overcoming friction
– Is a major problem for swimmers

• Overcoming gravity is less of a problem for


swimmers
– Than for animals that move on land or fly

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Locomotion on Land
• Walking, running, hopping, or crawling on land
– Requires an animal to support itself and move
against gravity

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• Diverse adaptations for traveling on land
– Have evolved in various vertebrates

Figure 49.36

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Flying
• Flight requires that wings develop enough lift
– To overcome the downward force of gravity

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Comparing Costs of Locomotion
•The energy cost of locomotion
–Depends on the mode of locomotion and the
environment
EXPERIMENT Physiologists typically determine an animal’s rate of energy use during locomotion by measuring
its oxygen consumption or carbon dioxide production while it swims in a water flume, runs on a treadmill, or flies in a
wind tunnel. For example, the trained parakeet shown below is wearing a plastic face mask connected to a tube that
collects the air the bird exhales as it flies.

RESULTS This graph compares the energy cost, in joules per kilogram of
body mass per meter traveled, for animals specialized for running, flying, and
swimming (1 J = 0.24 cal). Notice that both axes are plotted on logarithmic scales.

CONCLUSION For animals of a given Flying


body mass, swimming is the most energy- Running
Energy cost (J/Kg/m)

CONCLUSION 10 2

efficient and running the least energy-


efficient mode of locomotion. In any mode, 10
a small animal expends more energy per
kilogram of body mass than a large animal. 1

10–1 Swimming

10–3 1 103 106


Figure 49.37 Body mass(g)

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• Animals that are specialized for swimming
– Expend less energy per meter traveled than
equivalently sized animals specialized for
flying or running

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