Download chapter 49

• 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
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
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
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
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
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Annelids use their hydrostatic skeleton for
peristalsis
– A type of movement on land produced by
rhythmic waves of muscle contractions
(a) Body segments at the head and just in front
of the rear are short and thick (longitudinal
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.)
Longitudinal
muscle relaxed
(extended)
Bristles
(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.
Figure 49.25a–c
(c) The head segments are thick again and
anchored in their new positions. The rear
segments have released their hold on the
ground and have been pulled forward.
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Circular
muscle
contracted
Circular
muscle
relaxed
Longitudinal
muscle
contracted
Head
Exoskeletons
• An exoskeleton is a hard encasement
– Deposited on the surface of an animal
• Exoskeletons
– Are found in most molluscs and arthropods
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
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
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• 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
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• The human skeleton
key
Axial skeleton
Appendicular
skeleton
Skull
Examples
of joints
Head of
humerus
Scapula
1
Shoulder
girdle
Clavicle
Scapula
Sternum
Rib
Humerus
2
Vertebra
3
Radius
Ulna
Pelvic
girdle
1 Ball-and-socket joints, where the humerus contacts
the shoulder girdle and where the femur contacts the
pelvic girdle, enable us to rotate our arms and
legs and move them in several planes.
Humerus
Carpals
Phalanges
Ulna
Metacarpals
Femur
Patella
2 Hinge joints, such as between the humerus and
the head of the ulna, restrict movement to a single
plane.
Tibia
Fibula
Ulna
Figure 49.26
Tarsals
Metatarsals
Phalanges
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Radius
3 Pivot joints allow us to rotate our forearm at the
elbow and to move our head from side to side.
Physical Support on Land
• In addition to the skeleton
– Muscles and tendons help support large land
vertebrates
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Concept 49.6: Muscles move skeletal parts by
contracting
• The action of a muscle
– Is always to contract
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Skeletal muscles are attached to the skeleton
in antagonistic pairs
– With each member of the pair working against
each other
Human
Grasshopper
Extensor
muscle
relaxes
Biceps
contracts
Triceps
relaxes
Flexor
muscle
contracts
Forearm
flexes
Extensor
muscle
contracts
Biceps
relaxes
Forearm
extends
Figure 49.27
Triceps
contracts
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Tibia
flexes
Tibia
extends
Flexor
muscle
relaxes
Vertebrate Skeletal Muscle
• Vertebrate skeletal muscle
– Is characterized by a hierarchy of smaller and
smaller units
Muscle
Bundle of
muscle fibers
Nuclei
Single muscle fiber
(cell)
Plasma membrane
Myofibril
Z line
Light
band
animation
Dark band
Sarcomere
0.5 m
TEM
I band
A band
I band
M line
Thick
filaments
(myosin)
Figure 49.28
Thin
filaments
(actin)
Z line
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
H zone
Sarcomere
Z line
• A skeletal muscle consists of a bundle of long
fibers
– Running parallel to the length of the muscle
• A muscle fiber (muscle cell)
– Is itself a bundle of smaller myofibrils arranged
longitudinally
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• 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
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Skeletal muscle is also called striated muscle
– Because the regular arrangement of the
myofilaments creates a pattern of light and
dark bands
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
•
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
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
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
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• As a result of this sliding
– The I band and the H zone shrink
0.5 m
(a) Relaxed muscle fiber. In a relaxed muscle fiber, the I bands
and H zone are relatively wide.
(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.
Figure 49.29a–c
(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
the thick filaments contact the Z lines.
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Z
H
A
Sarcomere
• 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
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Myosin-actin interactions underlying muscle
fiber contraction
Thick filament
1 Starting here, the myosin head is
bound to ATP and is in its lowenergy confinguration.
Thin filaments
5 Binding of a new molecule of ATP releases the
myosin head from actin,
and a new cycle begins.
Thin filament
Myosin head (lowenergy configuration)
ATP
ATP
Thick
filament
Thin filament moves
toward center of sarcomere.
Figure 49.30
+
Cross-bridge
binding site
Actin
ADP
Myosin head (lowenergy configuration)
ADP
2 The myosin head hydrolyzes
ATP to ADP and inorganic
phosphate ( P I ) and is in its
high-energy configuration.
Pi
ADP
Pi
4 Releasing ADP and ( P i), myosin
relaxes to its low-energy configuration,
sliding the thin filament.
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Pi
Cross-bridge
Myosin head (highenergy configuration)
13 The myosin head binds to
actin, forming a crossbridge.
The Role of Calcium and Regulatory Proteins
• A skeletal muscle fiber contracts
– Only when stimulated by a motor neuron
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• When a muscle is at rest
– The myosin-binding sites on the thin filament
are blocked by the regulatory protein
tropomyosin
Tropomyosin
Actin
Figure 49.31a
Ca2+-binding sites
(a) Myosin-binding sites blocked
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Troponin complex
• 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+
Myosinbinding site
Figure 49.31b
(b) Myosin-binding sites exposed
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• 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
neuron axon
Mitochondrion
Synaptic
terminal
T tubule
Sarcoplasmic
reticulum
Ca2+ released
from sarcoplasmic
reticulum
Myofibril
Figure 49.32
Plasma membrane
of muscle fiber
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Sarcomere
• The synaptic terminal of the motor neuron
– Releases the neurotransmitter acetylcholine,
depolarizing the muscle and causing it to
produce an action potential
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• 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
terminal
of motor
neuron
1 Acetylcholine (ACh) released by synaptic terminal diffuses across synaptic
cleft and binds to receptor proteins on muscle fiber’s plasma membrane,
triggering an action potential in muscle fiber.
Synaptic cleft
ACh
2 Action potential is propagated along plasma
membrane and down
T tubules.
SR
3 Action potential
triggers Ca2+
release from sarcoplasmic reticulum
(SR).
Ca2
7 Tropomyosin blockage of myosinbinding sites is restored; contraction
ends, and muscle fiber relaxes.
Ca2
CYTOSOL
ADP
P2
PLASMA MEMBRANE
T TUBULE
4 Calcium ions bind to troponin;
troponin changes shape,
removing blocking action
of tropomyosin; myosin-binding
sites exposed.
2+
6 Cytosolic Ca is
removed by active
transport into
SR after action
potential ends.
Figure 49.33
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
5 Myosin cross-bridges alternately attach
to actin and detach, pulling actin
filaments toward center of sarcomere;
ATP powers sliding of filaments.
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
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• 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
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• 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
unit 1
Spinal cord
Motor
unit 2
Synaptic terminals
Nerve
Motor neuron
cell body
Motor neuron
axon
Muscle
Muscle fibers
Figure 49.34
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Tendon
• 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
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• A twitch
– Results from a single action potential in a
motor neuron
• More rapidly delivered action potentials
– Produce a graded contraction by summation
Tension
Tetanus
Summation of
two twitches
Single
twitch
Action
potential
Time
Pair of
action
potentials
Figure 49.35
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Series of action
potentials at
high frequency
• Tetanus is a state of smooth and sustained
contraction
– Produced when motor neurons deliver a volley
of action potentials
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
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
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Types of skeletal muscles
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
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
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• 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
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• 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
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
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
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Locomotion on Land
• Walking, running, hopping, or crawling on land
– Requires an animal to support itself and move
against gravity
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Diverse adaptations for traveling on land
– Have evolved in various vertebrates
Figure 49.36
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
Flying
• Flight requires that wings develop enough lift
– To overcome the downward force of gravity
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
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
Flying
Energy cost (J/Kg/m)
For animals of a given
body
mass, swimming is the most energyCONCLUSION
efficient and running the least energyefficient mode of locomotion. In any mode,
a small animal expends more energy per
kilogram of body mass than a large animal.
102
Running
10
1
Swimming
10–1
10–3
Figure 49.37
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
1
103
Body mass(g)
106