醫學研究所 Dr.

王志煜
Phone : 5092
E-mail :
jizyuhwang@cc.kmu.edu.tw

Chapter 50

Sensation and
Movement in
Animals

Lecture Presentations by
Nicole Tunbridge and
© 2018 Pearson Education Ltd. Kathleen Fitzpatrick
Concept 50.1:
Sensory receptors transduce stimulus energy
and transmit signals to the CNS
• All stimuli represent forms of energy
• Sensation involves converting energy into a change
in the membrane potential of sensory receptors
• When a stimulus’s input to the nervous system is
processed, a motor response may be generated
• This may involve a simple reflex or more elaborate
processing
© 2011 Pearson Education, Inc.
 Mole forages
along tunnel.
A simple response pathway:
foraging by a star-nosed mole

Mole
moves on.
Food absent

OR

Food present Mole bites.

Sensory input Integration Motor output

• Sensory pathways have four basic functions in
common
– Sensory reception
– Tranduction
– Transmission
– Perception (Integration)

© 2011 Pearson Education, Inc.

 Sensory Reception and Transduction

• Sensory receptors interact directly with stimuli,

both inside and outside the body
• Sensory transduction is the conversion of stimulus
energy into a change in the membrane potential of a
sensory receptor (= change cell activity)

• This change in membrane potential is called a

receptor potential

© 2011 Pearson Education, Inc.

Categories of sensory receptors
 Transmission
• Some sensory receptors are specialized neurons,
whereas others are specialized cells that regulate
neurons
• Sensory neurons produce action potentials and their
axons extend into the CNS
• The response of a sensory receptor varies with
intensity of stimuli
• If the receptor is a sensory neuron, a larger receptor
potential results in more frequent action potentials

© 2011 Pearson Education, Inc.

Coding of stimulus intensity by a single sensory receptor

Gentle pressure

Sensory receptor Low frequency of

action potentials per receptor

More pressure

High frequency of
action potentials per receptor
 Perception (Integration)
• Perceptions are the brain’s construction of stimuli
• Stimuli from different sensory receptors travel as
action potentials along dedicated neural pathways
• The brain distinguishes stimuli from different receptors
based on the area in the brain where the action
potentials arrive

© 2011 Pearson Education, Inc.

 Amplification and Adaptation

• The transduction of stimuli by sensory receptors is

subject to two types of modification
• Amplification is the strengthening of sensory signal
by neurons in transduction pathways
• Sensory adaptation is a decrease in
responsiveness to continued stimulation (e.g. heart
beat, clothing) because of receptor desensitization

© 2011 Pearson Education, Inc.

 Types of Sensory Receptors

Based on the nature of the stimuli to be transduced,

sensory receptors fall into five categories
– Mechanoreceptors
– Chemoreceptors
– Electromagnetic receptors
– Thermoreceptors
– Pain receptors

© 2011 Pearson Education, Inc.

 Mechanoreceptors
• Mechanoreceptors sense physical deformation
caused by stimuli such as pressure, touch, stretch,
motion, and sound
• They typically consist of ion channels linked to
structures that end outside the cell, such as “hairs”
(cilia)
• The mammalian sense of touch relies on
mechanoreceptors that are dendrites of sensory
neurons

© 2011 Pearson Education, Inc.

 Sensory receptors in human skin
Gentle pressure, vibration, Connective
and temperature tissue Pain
Hair

Epidermis

Dermis

Strong
pressure

Hypodermis

Nerve
Hair movement detected by naked dendrites
 Chemoreceptors
• General chemoreceptors transmit information about
the total solute concentration of a solution (e.g.
osmoreceptor)
• Specific chemoreceptors respond to individual kinds
of molecules (e.g. O2, CO2, glucose)
• When a stimulus molecule binds to a chemoreceptor,
the chemoreceptor becomes more or less permeable
to ions
• The antennae of the male silkworm moth (雄蠶蛾)
have very sensitive and specific chemoreceptors
© 2011 Pearson Education, Inc.
Chemoreceptors in an insect

0.1 mm
 Electromagnetic Receptors
• Electromagnetic receptors detect electromagnetic
energy such as visible light, electricity, and
magnetism
• The platypus has electroreceptors on its bill that can
detect the electric field generated by prey
• Many animals apparently migrate using the Earth’s
magnetic field to orient themselves

© 2011 Pearson Education, Inc.

Examples of electromagnetic reception and thermoreception
 Thermoreceptors
• Thermoreceptors detect heat and cold
• Certain snakes rely on thermoreceptors to detect
infrared radiation emitted by warm prey
• Jalapeno and cayenne peppers contain a substance
called capsaicin
• Receptors that respond to capsaicin respond to high
temperatures also, by opening a calcium channel
• Mammals have a variety of thermoreceptors, each
specific for a particular temperature range
© 2011 Pearson Education, Inc.
 Pain Receptors
• To detect stimuli that reflect the noxious (or harmful)
conditions
• In humans, pain receptors, or nociceptors, are a
class of naked dendrites in the epidermis
• They respond to excess heat, pressure, or chemicals
released from damaged or inflamed tissues
• Capsaicin receptor is a thermoreceptor and is also a
nociceptor

© 2011 Pearson Education, Inc.

 Concept 50.2:
The mechanoreceptors responsible for
hearing and equilibrium detect moving fluid
or settling particles
• Hearing and perception of body equilibrium are related
in most animals
• For both senses, settling particles or moving fluid are
detected by mechanoreceptors
 deflection (偏斜) of cell surface structures

© 2011 Pearson Education, Inc.

 Sensing Gravity in Invertebrates
• Most invertebrates maintain
equilibrium using
mechanoreceptors located in
organs called statocysts
• Statocysts contain
mechanoreceptors that detect
the movement of granules
called statoliths (耳石)
• Statoliths provide information
about the body position with
respect to gravity

© 2011 Pearson Education, Inc.

 Sensing Sound in Invertebrates
• Many insects have body hairs that vibrate in response to sound
waves
• Many also detect sound with localized ears consisting of a
tympanic membrane (鼓膜) stretched over an internal air
chamber

Tympanic
membrane

1 mm
 Hearing and Equilibrium in Mammals

• In most terrestrial vertebrates, sensory organs

for hearing and equilibrium are closely
associated in the ear

© 2011 Pearson Education, Inc.

 Outer ear Middle ear Inner ear

Skull Stapes Semicircular

bone Cochlear Bone
Incus canals
duct Auditory
Malleus Auditory nerve nerve
to brain Vestibular
canal

Tympanic
canal
Cochlea Organ
Oval Eustachian of Corti
Auditory window
Pinna canal tube
Tympanic Round
membrane window Tectorial membrane
1 m

Hair cells Axons of

To auditory
Basilar sensory
Bundled hairs projecting from a hair cell nerve
membrane neurons
 Hearing
• Vibrating objects create pressure waves in the air that
cause the tympanic membrane to vibrate
• The three bones of the middle ear transmit the
vibrations of moving air to the oval window on the
cochlea (耳蝸)
• These vibrations create pressure waves in the fluid in
the cochlea that travel through the vestibular canal
(前庭腔)
• The fluid waves dissipate when they strike the round
window at the end of the tympanic canal (鼓腔)
© 2011 Pearson Education, Inc.
 Hearing
• Fluid pressure waves push down on the cochlear duct
and basilar membrane
• The basilar membrane and the attached hair cells
vibrate up and down
• This bending of hair cells depolarizes the membranes
of mechanoreceptors and sends action potentials to
the brain via the auditory nerve

© 2011 Pearson Education, Inc.

 Sensory reception by hair cells
“Hairs” of
hair cell
Neurotrans- More
mitter at neuro- Less
synapse trans- neuro-
mitter trans-
mitter
Sensory 50 50 Receptor potential 50
neuron
potential (mV)

potential (mV)

potential (mV)
70 70 70
Membrane

Membrane

Membrane
Action potentials
Signal

Signal
Signal

0 0 0

70 70 70

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

(a) No bending of hairs (b) Bending of hairs in one direction (c) Bending of hairs in other direction

毛束擺盪方向影響神經傳遞物的釋放量
• The ear conveys information about
– Volume (音量), the amplitude of the sound wave
– Pitch (音高), the frequency of the sound wave
• The cochlea can distinguish pitch because the basilar
membrane is not uniform along its length
A B C
Axons of Point C 3 6,000 Hz
sensory neurons

Relative motion of basilar membrane

Apex
Oval
window
B 0
Stapes Vestibular
C canal 3 1,000 Hz
A

Cochlea
0
Point B
Tympanic 3 100 Hz
membrane Basilar
membrane

Base
Round Tympanic 0
window canal 0 10 20 30
Distance from oval window (mm)
Point A
(a) (b)
 Equilibrium
• Several organs of the inner ear detect body
movement, position, and balance
– The utricle (橢圓囊) and saccule (球囊) of vestibule
contain granules called otoliths (耳石) that allow us to
perceive position relative to gravity or linear
movement
– Three semicircular canals contain fluid and can detect
angular movement in any direction

© 2011 Pearson Education, Inc.

 Organs of equilibrium in the inner ear
Semicircular PERILYMPH
canals Cupula
Fluid
flow
Vestibular
nerve

Hairs
Hair
cell
Vestibule Nerve
fibers
Utricle
Body movement
Saccule
 Concept 50.3:
Visual receptors on diverse animals depend
on light-absorbing pigments

© 2011 Pearson Education, Inc.

 Evolution of Visual Perception
• Animals use a diverse set of organs for vision, but
the underlying mechanism for capturing light is the
same.
• Light detectors in the animal kingdom range from
simple clusters of cells that detect direction and
intensity of light to complex organs that form images
• Light detectors all contain photoreceptors, cells
that contain light-absorbing pigment molecules

© 2011 Pearson Education, Inc.

 Light-Detecting Organs
• Most invertebrates have a light-detecting organ
• One of the simplest light-detecting organs is that of
planarians
• A pair of ocelli (單眼) called eyespots are located
near the head
• These allow planarians to move away from light and
seek shaded locations

© 2011 Pearson Education, Inc.

Ocelli and orientation behavior of a planarian
LIGHT

DARK
(a)

Light

Photoreceptor
Ocellus Nerve to
Visual brain
pigment
Screening
(b) Ocellus pigment
 Compound Eyes
• Insects and crustaceans
have compound eyes,
which consist of up to
several thousand light
detectors called ommatidia
(小眼)
• Compound eyes are very
effective at detecting
movement
• Insects have excellent color
vision, and some can see
into the ultraviolet range
© 2011 Pearson Education, Inc.
 Single-Lens Eyes
• Single-lens eyes are found in some jellies,
polychaete worms, spiders, and many molluscs
• They work on a camera-like principle: the iris(虹膜)
changes the diameter of the pupil to control how
much light enters
• The eyes of all vertebrates has a single lens
• In vertebrates the eye detects color and light, but the
brain assembles the information and perceives the
image
© 2011 Pearson Education, Inc.
 The Vertebrate Visual System
Choroid Retina
Sclera
Retina Photoreceptors
Suspensory Neurons
Fovea Rod Cone
ligament
Cornea
Iris
Optic
nerve
Pupil

Aqueous
humor
Lens Central
artery and
vein of
Vitreous humor Optic disk the retina
Optic Amacrine Horizontal cell
nerve cell Pigmented
fibers Ganglion Bipolar epithelium
cell cell

• Humans have two main types of photoreceptor cells

– Rods are more sensitive to light but do not distinguish colors
– Cones provide color vision
Rod CYTOSOL

Synaptic Cell Outer Disks Retinal: cis isomer

terminal body segment
Light Enzymes
Cone

Rod
Retinal: trans isomer
Cone
Retinal
Rhodopsin
Opsin
INSIDE OF DISK
 Sensory Transduction in the Eye
• Transduction of visual information to the nervous
system begins with the light-induced conversion of
cis-retinal to trans-retinal
• trans-retinal activates rhodopsin (視紫質), which
activates a G protein, eventually leading to hydrolysis
of cyclic GMP
• When cyclic GMP breaks down, Na channels close
• This hyperpolarizes the cell
• When enzymes convert retinal back to the cis form,
returning rhodopsin to its inactive state, the signal
transduction pathway normally shuts off
© 2011 Pearson Education, Inc.
 Response of a photoreceptor cell to light
DARK Inactive INSIDE OF DISK Disk Plasma
rhodopsin Membrane potential (mV)
membrane membrane
Rod cell across plasma membrane
0
Depolarized
state
–40
cGMP
cGMP Na+
Transducin cGMP –70
Inactive cGMP Time
cGMP
CYTOSOL phosphodiesterase

LIGHT Active INSIDE OF DISK Disk Plasma Membrane potential (mV)

rhodopsin membrane membrane across plasma membrane
0
Light strikes
Synaptic retina
terminals Na+
cGMP –40
Hyper-
GMP Transducin –70
GMP polarization
GMP GMP Activated GMP
CYTOSOL Time
phosphodiesterase
 Dark Responses Light Responses

Rhodopsin inactive Rhodopsin active

Na channels open Na channels closed

Rod depolarized Rod hyperpolarized

No glutamate
Glutamate released released

Bipolar cell either Bipolar cell either

depolarized or hyperpolarized or
hyperpolarized, depolarized,
depending on depending on
glutamate receptors glutamate receptors
Processing of Visual Information in the Retina
• Three other types of neurons contribute to information
processing in the retina
– Ganglion cells transmit signals from bipolar cells to the
brain
– Horizontal and amacrine cells help integrate visual
information before it is sent to the brain
• Interaction among different cells results in lateral
inhibition, enhanced contrast in the image
• Lateral inhibition is when a rod or cone stimulates a
horizontal cell and the horizontal cell then inhibits
more distant photoreceptors and bipolar cells
© 2011 Pearson Education, Inc.
Processing of Visual Information in the Brain

• The optic nerves meet at the optic chiasm (視交叉)

near the cerebral cortex
• Most ganglion cell axons lead to the lateral
geniculate nuclei (外側膝狀體核)
• The lateral geniculate nuclei relay information to the
primary visual cortex in the cerebrum

© 2011 Pearson Education, Inc.

 Sensations from the left visual field of both eyes are transmitted
to the right side of the brain
 Sensations from the right visual field of both eyes are
transmitted to the left side of the brain
 Color Vision
• Mammals that are nocturnal usually have a high
proportion of rods in the retina
• In humans, perception of color is based on three types
of cones, each with a different visual pigment: red,
green, or blue
• The three visual pigments, called photopsins (光視質),
are formed when retinal binds to three distinct opsin
proteins (視蛋白)
• Abnormal color vision results from alterations in the
genes encoding one or more photopsin proteins

© 2011 Pearson Education, Inc.

 The Visual Field
• The brain processes visual information and controls
what information is captured
• Focusing occurs by changing the shape of the lens
• The fovea (中央窩) is the center of the visual field
and contains no rods, but a high density of cones

© 2011 Pearson Education, Inc.

 Focusing in the mammalian eye
(a) Near vision (accommodation) (b) Distance vision

Ciliary muscles Ciliary muscles

contract, pulling relax, and border
border of choroid Choroid of choroid moves
toward lens. away from lens.
Retina
Suspensory
Suspensory ligaments pull
ligaments against lens.
relax.

Lens becomes
thicker and
rounder, focusing Lens becomes
on nearby objects. flatter, focusing on
distant objects.
Concept 50.4:
The senses of taste and smell rely on similar
sets of sensory receptors
• In terrestrial animals
– Gustation (= taste) is dependent on the detection of
chemicals called tastants
– Olfaction (= smell) is dependent on the detection of
chemicals called odorants
• In aquatic animals, there is no distinction between
taste and smell
• Taste receptors of insects are in sensory hairs called
sensilla (感器), located on feet and in mouthparts

© 2011 Pearson Education, Inc.

 Taste in Mammals
• In humans, receptor cells for taste are modified
epithelial cells organized into taste buds
• There are five taste perceptions: sweet, sour, salty,
bitter, and umami (甘味) (elicited by glutamate)
• Researchers have identified receptors for all five
tastes
• Researchers show that an individual taste cell
expresses a single receptor type and detects one of
the five tastes
• Any region with taste buds can detect any of the five
types of taste
© 2011 Pearson Education, Inc.
 Papilla

Most taste buds are

Papillae
associated with projections Taste
buds
called papillae
(a) Tongue

Key Taste bud Saliva

Sweet
Salty
Taste
Sour pore
Bitter
Umami

Sensory
neuron Sensory Food
receptor cells molecules
(b) Taste buds
 Taste in Mammals
 Receptor cells for taste in mammals are modified
epithelial cells organized into taste buds, located in
several areas of the tongue and mouth
 Taste receptors are of three types
 The sensations of sweet, umami, and bitter require
specific G protein-coupled receptors (GPCRs)
 The receptor for sour belongs to the TRP family and is
similar to the capsaicin and other thermoreceptor
proteins
 The taste receptor for salt is a sodium channel
 Smell in Humans
• Olfactory receptor cells are neurons that line the
upper portion of the nasal cavity
• Binding of odorant molecules to receptors triggers a
signal transduction pathway, sending action
potentials to the brain
• Humans can distinguish thousands of different odors
• Although receptors and brain pathways for taste and
smell are independent, the two senses do interact

© 2011 Pearson Education, Inc.

 Brain
potentials

Olfactory
bulb
Odorants
Nasal cavity Bone

Receptors Epithelial
for different cell
odorants
Chemo-
receptor
Plasma Cilia
membrane

Odorants Mucus
Concept 50.5:
The physical interaction of protein filaments
is required for muscle function
• Muscle activity is a response to input from the nervous
system
• The action of a muscle is always to contract; extension
is passive

© 2011 Pearson Education, Inc.

 Vertebrate Skeletal Muscle
• Vertebrate skeletal muscle moves bones and the
body, and is characterized by a hierarchy of smaller
and smaller units
• A skeletal muscle consists of a bundle of long fibers,
each a single cell, running parallel to the length of
the muscle
• Each muscle fiber is itself a bundle of smaller
myofibrils (肌原纖維) arranged longitudinally

© 2011 Pearson Education, Inc.

 Muscle

Bundle of
muscle fibers
Nuclei
Single muscle fiber (cell)
Plasma membrane

Myofibril
Z lines

Sarcomere
 Vertebrate Skeletal Muscle
• The myofibrils are composed to two kinds of
myofilaments
– Thin filaments consist of two strands of actin and
two strands of a regulatory protein
– Thick filaments are staggered arrays of myosin
molecules
• Skeletal muscle is also called striated muscle
because the regular arrangement of myofilaments
creates a pattern of light and dark bands
• The functional unit of a muscle is called a sarcomere
and is bordered by Z lines
© 2011 Pearson Education, Inc.
 Z lines

Sarcomere

TEM
Thick M line 0.5 m
filaments
(myosin)

Thin
filaments
(actin)
Z line Z line
Sarcomere
 The Sliding-Filament Model of Muscle
Contraction
• According to the sliding-filament model, filaments
slide past each other longitudinally, producing more
overlap between thin and thick filaments
• The sliding of filaments relies on interaction between
actin and myosin

© 2011 Pearson Education, Inc.

 The Sliding-Filament Model of Muscle
Contraction
• 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
• Muscle contraction requires repeated cycles of
binding and release
• Glycolysis and aerobic respiration generate the ATP
needed to sustain muscle contraction

© 2011 Pearson Education, Inc.

 Figure 50.28
Thin
filaments

Thick filament 1

Thin filament

ATP Myosin head (low-

energy configuration)
ATP 2
Thick
5 filament
Myosin-
Thin filament moves binding sites
toward center of sarcomere. Actin

Myosin head (low- ADP Myosin head (high-

energy configuration) Pi
energy configuration

ADP 3
ADP Pi Pi Cross-bridge
4
The Role of Calcium and Regulatory Proteins
• The regulatory protein
tropomyosin and the
troponin complex, a
set of additional
proteins, bind to actin
strands on thin
filaments when a
muscle fiber is at rest
• This prevents actin and
myosin from interacting

© 2011 Pearson Education, Inc.

 The Role of Calcium and Regulatory Proteins
• For a muscle fiber to contract, myosin-binding sites
must be uncovered
• This occurs when calcium ions (Ca2+) bind to the
troponin complex and expose the myosin-binding sites
• Contraction occurs when the concentration of Ca2+ is
high; muscle fiber contraction stops when the
concentration of Ca2+ is low

© 2011 Pearson Education, Inc.

 The Sliding-Filament Model of Muscle
Contraction

https://www.youtube.com/watch?v=7O_ZHyPeIIA
The Role of Calcium and Regulatory Proteins
• The stimulus leading to contraction of a muscle fiber
is an action potential in a motor neuron that makes a
synapse with the muscle fiber
• The synaptic terminal of the motor neuron releases
the neurotransmitter acetylcholine
• Acetylcholine depolarizes the muscle, causing it to
produce an action potential

© 2011 Pearson Education, Inc.

The Role of Calcium and Regulatory Proteins
• Action potentials travel to the interior of the muscle
fiber along transverse (T) tubules
• The action potential along T tubules causes the
sarcoplasmic reticulum (SR, 肌漿網) to release Ca2+
• The Ca2+ binds to the troponin complex on the thin
filaments
• This binding exposes myosin-binding sites and allows
the cross-bridge cycle to proceed

© 2011 Pearson Education, Inc.

 1
Synaptic terminal of motor neuron
Synaptic cleft T tubule Plasma membrane

2 Sarcoplasmic
ACh reticulum (SR)

Ca2 3
Ca2 pump

ATP

6 4
CYTOSOL
Ca2
7

5
The Role of Calcium and Regulatory Proteins

• When motor neuron input stops, the muscle cell

relaxes
• ATP-dependent transport proteins in the SR pump
Ca2+ out of the cytosol
• Regulatory proteins bound to thin filaments shift back
to the myosin-binding sites

© 2011 Pearson Education, Inc.

 Nervous Control of Muscle Tension

• In vertebrates, each motor neuron may synapse with

multiple muscle fibers, although each fiber is
controlled by only one motor neuron
• A motor unit consists of a single motor neuron and
all the muscle fibers it controls

© 2011 Pearson Education, Inc.

 Spinal cord Motor Motor
unit 1 unit 2
Synaptic
terminals

Nerve

Motor neuron
cell body
Motor neuron
axon

Muscle

Muscle fibers

Tendon
Concept 50.6:
Skeletal systems transform muscle
contraction into locomotion

• The skeleton provides a rigid structure to which

muscles attach
• Skeletons function in support, protection, and
movement

© 2011 Pearson Education, Inc.

 Types of Skeletal Systems

• The three main types of skeletons are

– Hydrostatic skeletons (lack hard parts)
– Exoskeletons (external hard parts)
– Endoskeletons (internal hard parts)

© 2011 Pearson Education, Inc.

 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
• Annelids use their hydrostatic skeleton for
peristalsis (蠕動), a type of movement on land
produced by rhythmic waves of muscle contractions

© 2011 Pearson Education, Inc.

Longitudinal Circular Circular Longitudinal
muscle relaxed muscle muscle muscle
(extended) contracted relaxed contracted

Bristles
(剛毛) Head end

Head end

Head end

3
 Exoskeletons
• An exoskeleton is a hard encasement deposited on
the surface of an animal
• Exoskeletons are found in most molluscs and
arthropods
• Arthropods have a jointed exoskeleton called a cuticle
(角質), which can be both strong and flexible
• The polysaccharide chitin (多醣幾丁質) is often found in
arthropod cuticle

© 2011 Pearson Education, Inc.

 Endoskeletons
• An endoskeleton consists of a hard internal
skeleton, buried in soft tissue
• Endoskeletons are found in organisms ranging from
sponges to mammals
• A mammalian skeleton has more than 200 bones
• Some bones are fused; others are connected at
joints by ligaments that allow freedom of movement

© 2011 Pearson Education, Inc.

 Types
Shoulder Skull of joints
Clavicle
girdle Ball-and-socket
Scapula joint
Sternum
Rib
Humerus Hinge joint
Vertebra Pivot joint
Radius
Ulna
Pelvic girdle
Carpals

Phalanges
Metacarpals
Femur
Patella
Tibia
Fibula

Tarsals
Metatarsals
Phalanges 206
 Types of Locomotion
• Most animals are capable of locomotion, or active
travel from place to place
• In locomotion, energy is expended to overcome
friction and gravity

Locomotion on Land
 Walking, running, hopping, or crawling on land
requires an animal to support itself and move against
gravity
 Diverse adaptations for locomotion on land have
evolved in vertebrates
© 2011 Pearson Education, Inc.
 Swimming
• In water, drag (friction) is a bigger problem than gravity
• Fast swimmers usually have a sleek, fusiform shape
to minimize friction
• Animals swim in diverse ways
– Paddling with their legs as oars
– Jet propulsion
– Undulating their body and tail from side to side, or
up and down

© 2011 Pearson Education, Inc.

 Flying
• Active flight requires that wings develop enough lift to
overcome the downward force of gravity
• Many flying animals have adaptations that reduce
body mass
– For example, birds lack teeth and a urinary bladder
(膀胱), as well as large bones with air-filled regions

© 2011 Pearson Education, Inc.

• The energy cost of locomotion can be estimated by
the rate of oxygen consumption or carbon dioxide
production
Figure 50.39
Data from the Experiments

Flying
Energy cost [cal/(kg·m)]

Running
102
(log scale)

10

1
Swimming
10–1

10–3 1 103 106

Body mass (g) (log scale)
Data from K. Schmidt-Nielsen, Locomotion: Energy cost of swimming, flying, and running,
Science 177:222–228 (1972). Reprinted with permission from AAAS.
 Energy Costs of Locomotion
• Animals specialized for swimming expend less energy per
meter traveled than equivalently sized animals specialized for
flying or running
RESULTS
Flying Running
Energy cost (cal/kg m)

102
(log scale)

10

1
Swimming
101

103 1 103 106

Body mass (g) (log scale)
 Energy Costs of Locomotion
• Large animals travel more efficiently than smaller animals
adapted to the same mode of transport (locomotion)
RESULTS
Flying Running
Energy cost (cal/kg m)

102
(log scale)

10 較耗能

1 較不耗能
Swimming
101
小型動物 大型動物

103 1 103 106

Body mass (g) (log scale)