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Chapter 10
Muscle Tissue
10-1 An Introduction to Muscle Tissue
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Learning Outcomes
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10-1 Specify the functions of skeletal muscle tissue.
10-2 Describe the organization of muscle at the tissue level.
10-3 Explain the characteristics of skeletal muscle fibers, and
identify the structural components of a sarcomere.
10-4 Identify the components of the neuromuscular junction, and
summarize the events involved in the neural control of
skeletal muscle contraction and relaxation.
10-1 An Introduction to Muscle Tissue
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Learning Outcomes
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10-5 Describe the mechanism responsible for tension
production in a muscle fiber, and compare the different
types of muscle contraction.
10-6 Describe the mechanisms by which muscle fibers obtain
the energy to power contractions.
10-7 Relate the types of muscle fibers to muscle performance,
and distinguish between aerobic and anaerobic endurance.
10-1 An Introduction to Muscle Tissue
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Learning Outcomes
© 2012 Pearson Education, Inc.
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10-8 Identify the structural and functional differences between
skeletal muscle fibers and cardiac muscle cells.
10-9 Identify the structural and functional differences between
skeletal muscle fibers and smooth muscle cells, and
discuss the roles of smooth muscle tissue in systems
throughout the body.
An Introduction to Muscle Tissue
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Muscle Tissue
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A primary tissue type, divided into:
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Skeletal muscle tissue
Cardiac muscle tissue
Smooth muscle tissue
10-1 Functions of Skeletal Muscle Tissue
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Skeletal Muscles
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Are attached to the skeletal system
Allow us to move
The muscular system
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Includes only skeletal muscles
10-1 Functions of Skeletal Muscle Tissue
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Six Functions of Skeletal Muscle Tissue
© 2012 Pearson Education, Inc.
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Produce skeletal movement
Maintain posture and body position
Support soft tissues
Guard entrances and exits
Maintain body temperature
Store nutrient reserves
10-2 Organization of Muscle
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Skeletal Muscle
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Muscle tissue (muscle cells or fibers)
Connective tissues
Nerves
Blood vessels
10-2 Organization of Muscle
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Organization of Connective Tissues
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Muscles have three layers of connective tissues
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Epimysium
Perimysium
Endomysium
10-2 Organization of Muscle
© 2012 Pearson Education, Inc.
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Epimysium
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Exterior collagen layer
Connected to deep fascia
Separates muscle from surrounding tissues
10-2 Organization of Muscle
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Perimysium
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Surrounds muscle fiber bundles (fascicles)
Contains blood vessel and nerve supply to fascicles
10-2 Organization of Muscle
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Endomysium
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Surrounds individual muscle cells (muscle fibers)
Contains capillaries and nerve fibers contacting muscle cells
Contains myosatellite cells (stem cells) that repair damage
10-2 Organization of Muscle
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Organization of Connective Tissues
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Muscle Attachments
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Endomysium, perimysium, and epimysium come together:
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At ends of muscles
To form connective tissue attachment to bone matrix
© 2012 Pearson Education, Inc.
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I.e., tendon (bundle) or aponeurosis (sheet)
10-2 Organization of Muscle
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Blood Vessels and Nerves
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Muscles have extensive vascular systems that:
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Supply large amounts of oxygen
Supply nutrients
Carry away wastes
Skeletal muscles are voluntary muscles, controlled by nerves
of the central nervous system (brain and spinal cord)
10-3 Characteristics of Skeletal Muscle Fibers
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Skeletal Muscle Cells
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Are very long
Develop through fusion of mesodermal cells (myoblasts)
Become very large
Contain hundreds of nuclei
10-3 Characteristics of Skeletal Muscle Fibers
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The Sarcolemma and Transverse Tubules
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The sarcolemma
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The cell membrane of a muscle fiber (cell)
Surrounds the sarcoplasm (cytoplasm of muscle fiber)
© 2012 Pearson Education, Inc.
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A change in transmembrane potential begins contractions
10-3 Characteristics of Skeletal Muscle Fibers
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The Sarcolemma and Transverse Tubules
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Transverse tubules (T tubules)
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Transmit action potential through cell
Allow entire muscle fiber to contract simultaneously
Have same properties as sarcolemma
10-3 Characteristics of Skeletal Muscle Fibers
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Myofibrils
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Lengthwise subdivisions within muscle fiber
Made up of bundles of protein filaments (myofilaments)
Myofilaments are responsible for muscle contraction
Types of myofilaments:
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Thin filaments
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Made of the protein actin
Thick filaments
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Made of the protein myosin
10-3 Characteristics of Skeletal Muscle Fibers
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The Sarcoplasmic Reticulum (SR)
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A membranous structure surrounding each myofibril
© 2012 Pearson Education, Inc.
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Helps transmit action potential to myofibril
Similar in structure to smooth endoplasmic reticulum
Forms chambers (terminal cisternae) attached to T tubules
10-3 Characteristics of Skeletal Muscle Fibers
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The Sarcoplasmic Reticulum (SR)
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Triad
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Is formed by one T tubule and two terminal cisternae
Cisternae
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Concentrate Ca2+ (via ion pumps)
Release Ca2+ into sarcomeres to begin muscle contraction
10-3 Structural Components of a Sarcomere
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Sarcomeres
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The contractile units of muscle
Structural units of myofibrils
Form visible patterns within myofibrils
A striped or striated pattern within myofibrils
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Alternating dark, thick filaments (A bands) and light, thin
filaments (I bands)
10-3 Structural Components of a Sarcomere
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Sarcomeres
© 2012 Pearson Education, Inc.
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The A Band
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M line
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The center of the A band
At midline of sarcomere
The H Band
The area around the M line
Has thick filaments but no thin filaments
Zone of overlap
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The densest, darkest area on a light micrograph
Where thick and thin filaments overlap
10-3 Structural Components of a Sarcomere
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Sarcomeres
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The I Band
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Z lines
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The centers of the I bands
At two ends of sarcomere
Titin
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Are strands of protein
Reach from tips of thick filaments to the Z line
Stabilize the filaments
10-3 Structural Components of a Sarcomere
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Thin Filaments
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F-actin (filamentous actin)
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Is two twisted rows of globular G-actin
© 2012 Pearson Education, Inc.
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The active sites on G-actin strands bind to myosin
Nebulin
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Holds F-actin strands together
10-3 Structural Components of a Sarcomere
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Thin Filaments
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Tropomyosin
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Is a double strand
Prevents actin–myosin interaction
Troponin
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A globular protein
Binds tropomyosin to G-actin
Controlled by Ca2+
10-3 Structural Components of a Sarcomere
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Initiating Contraction
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Ca2+ binds to receptor on troponin molecule
Troponin–tropomyosin complex changes
Exposes active site of F-actin
10-3 Structural Components of a Sarcomere
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Thick Filaments
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Contain about 300 twisted myosin subunits
© 2012 Pearson Education, Inc.
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Contain titin strands that recoil after stretching
The mysosin molecule
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Tail
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Binds to other myosin molecules
Head
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Made of two globular protein subunits
Reaches the nearest thin filament
10-3 Structural Components of a Sarcomere
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Myosin Action
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During contraction, myosin heads:
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Interact with actin filaments, forming cross-bridges
Pivot, producing motion
10-3 Structural Components of a Sarcomere
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Sliding Filaments and Muscle Contraction
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Sliding filament theory
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Thin filaments of sarcomere slide toward M line, alongside
thick filaments
The width of A zone stays the same
Z lines move closer together
10-3 Structural Components of a Sarcomere
© 2012 Pearson Education, Inc.
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Skeletal Muscle Contraction
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The process of contraction
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Neural stimulation of sarcolemma
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Causes excitation–contraction coupling
Muscle fiber contraction
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Interaction of thick and thin filaments
Tension production
10-4 Components of the Neuromuscular Junction
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The Control of Skeletal Muscle Activity
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The neuromuscular junction (NMJ)
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Special intercellular connection between the nervous system
and skeletal muscle fiber
Controls calcium ion release into the sarcoplasm
10-4 Components of the Neuromuscular Junction
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Excitation–Contraction Coupling
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Action potential reaches a triad
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Releasing Ca2+
Triggering contraction
Requires myosin heads to be in “cocked” position
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Loaded by ATP energy
© 2012 Pearson Education, Inc.
10-4 Skeletal Muscle Contraction
• The Contraction Cycle
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Contraction Cycle Begins
Active-Site Exposure
Cross-Bridge Formation
Myosin Head Pivoting
Cross-Bridge Detachment
Myosin Reactivation
10-4 Skeletal Muscle Contraction
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Fiber Shortening
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As sarcomeres shorten, muscle pulls together, producing
tension
Muscle shortening can occur at both ends of the muscle, or at
only one end of the muscle
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This depends on the way the muscle is attached at the ends
10-4 Skeletal Muscle Relaxation
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Relaxation
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Contraction Duration
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Depends on:
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Duration of neural stimulus
© 2012 Pearson Education, Inc.
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Number of free calcium ions in sarcoplasm
Availability of ATP
10-4 Skeletal Muscle Relaxation
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Relaxation
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Ca2+ concentrations fall
Ca2+ detaches from troponin
Active sites are re-covered by tropomyosin
Rigor Mortis
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A fixed muscular contraction after death
Caused when:
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Ion pumps cease to function; ran out of ATP
Calcium builds up in the sarcoplasm
10-4 Skeletal Muscle Contraction and Relaxation
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Summary
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Skeletal muscle fibers shorten as thin filaments slide between
thick filaments
Free Ca2+ in the sarcoplasm triggers contraction
SR releases Ca2+ when a motor neuron stimulates the
muscle fiber
Contraction is an active process
© 2012 Pearson Education, Inc.
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Relaxation and return to resting length are passive
10-5 Tension Production and Contraction Types
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Tension Production by Muscles Fibers
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As a whole, a muscle fiber is either contracted or relaxed
Depends on:
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The number of pivoting cross-bridges
The fiber’s resting length at the time of stimulation
The frequency of stimulation
10-5 Tension Production and Contraction Types
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Tension Production by Muscles Fibers
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Length–Tension Relationships
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Number of pivoting cross-bridges depends on:
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Amount of overlap between thick and thin fibers
Optimum overlap produces greatest amount of tension
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Too much or too little reduces efficiency
Normal resting sarcomere length
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Is 75% to 130% of optimal length
10-5 Tension Production and Contraction Types
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Tension Production by Muscles Fibers
© 2012 Pearson Education, Inc.
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The Frequency of Stimulation
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A single neural stimulation produces:
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A single contraction or twitch
Which lasts about 7–100 msec.
Sustained muscular contractions
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Require many repeated stimuli
10-5 Tension Production and Contraction Types
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Tension Production by Muscles Fibers
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Twitches
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Latent period
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The action potential moves through sarcolemma
Causing Ca2+ release
Contraction phase
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Calcium ions bind
Tension builds to peak
Relaxation phase
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Ca2+ levels fall
Active sites are covered and tension falls to resting levels
10-5 Tension Production and Contraction Types
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Tension Production by Muscles Fibers
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Treppe
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A stair-step increase in twitch tension
Repeated stimulations immediately after relaxation phase
© 2012 Pearson Education, Inc.
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Stimulus frequency <50/second
Causes a series of contractions with increasing tension
10-5 Tension Production and Contraction Types
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Tension Production by Muscles Fibers
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Wave summation
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Increasing tension or summation of twitches
Repeated stimulations before the end of relaxation phase
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Stimulus frequency >50/second
Causes increasing tension or summation of twitches
10-5 Tension Production and Contraction Types
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Tension Production by Muscles Fibers
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Incomplete tetanus
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Twitches reach maximum tension
If rapid stimulation continues and muscle is not allowed to
relax, twitches reach maximum level of tension
Complete tetanus
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If stimulation frequency is high enough, muscle never begins
to relax, and is in continuous contraction
10-5 Tension Production and Contraction Types
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Tension Production by Skeletal Muscles
© 2012 Pearson Education, Inc.
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Depends on:
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Internal tension produced by muscle fibers
External tension exerted by muscle fibers on elastic
extracellular fibers
Total number of muscle fibers stimulated
10-5 Tension Production and Contraction Types
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Motor Units and Tension Production
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Motor units in a skeletal muscle:
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Contain hundreds of muscle fibers
That contract at the same time
Controlled by a single motor neuron
10-5 Tension Production and Contraction Types
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Motor Units and Tension Production
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Recruitment (multiple motor unit summation)
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In a whole muscle or group of muscles, smooth motion and
increasing tension are produced by slowly increasing the
size or number of motor units stimulated
Maximum tension
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Achieved when all motor units reach tetanus
Can be sustained only a very short time
© 2012 Pearson Education, Inc.
10-5 Tension Production and Contraction Types
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Motor Units and Tension Production
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Sustained tension
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Less than maximum tension
Allows motor units rest in rotation
Muscle tone
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The normal tension and firmness of a muscle at rest
Muscle units actively maintain body position, without motion
Increasing muscle tone increases metabolic energy used,
even at rest
10-5 Tension Production and Contraction Types
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Motor Units and Tension Production
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Contraction are classified based on pattern of tension
production
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Isotonic contraction
Isometric contraction
10-5 Tension Production and Contraction Types
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Isotonic Contraction
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Skeletal muscle changes length
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Resulting in motion
If muscle tension > load (resistance):
© 2012 Pearson Education, Inc.
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Muscle shortens (concentric contraction)
If muscle tension < load (resistance):
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Muscle lengthens (eccentric contraction)
10-5 Tension Production and Contraction Types
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Isometric Contraction
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Skeletal muscle develops tension, but is prevented from
changing length
iso- = same, metric = measure
10-5 Tension Production and Contraction Types
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Load and Speed of Contraction
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Are inversely related
The heavier the load (resistance) on a muscle
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The longer it takes for shortening to begin
And the less the muscle will shorten
10-5 Tension Production and Contraction Types
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Muscle Relaxation and the Return to Resting Length
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Elastic Forces
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The pull of elastic elements (tendons and ligaments)
Expands the sarcomeres to resting length
Opposing Muscle Contractions
© 2012 Pearson Education, Inc.
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Reverse the direction of the original motion
Are the work of opposing skeletal muscle pairs
10-5 Tension Production and Contraction Types
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Muscle Relaxation and the Return to Resting Length
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Gravity
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Can take the place of opposing muscle contraction to return
a muscle to its resting state
10-6 Energy to Power Contractions
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ATP Provides Energy For Muscle Contraction
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Sustained muscle contraction uses a lot of ATP energy
Muscles store enough energy to start contraction
Muscle fibers must manufacture more ATP as needed
10-6 Energy to Power Contractions
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ATP and CP Reserves
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Adenosine triphosphate (ATP)
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The active energy molecule
Creatine phosphate (CP)
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The storage molecule for excess ATP energy in resting muscle
Energy recharges ADP to ATP
© 2012 Pearson Education, Inc.
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Using the enzyme creatine kinase (CK)
When CP is used up, other mechanisms generate ATP
10-6 Energy to Power Contractions
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ATP Generation
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Cells produce ATP in two ways
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Aerobic metabolism of fatty acids in the mitochondria
Anaerobic glycolysis in the cytoplasm
10-6 Energy to Power Contractions
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Aerobic Metabolism
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Is the primary energy source of resting muscles
Breaks down fatty acids
Produces 34 ATP molecules per glucose molecule
Glycolysis
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Is the primary energy source for peak muscular activity
Produces two ATP molecules per molecule of glucose
Breaks down glucose from glycogen stored in skeletal muscles
10-6 Energy to Power Contractions
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Energy Use and the Level of Muscular Activity
© 2012 Pearson Education, Inc.
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Skeletal muscles at rest metabolize fatty acids and store
glycogen
During light activity, muscles generate ATP through
anaerobic breakdown of carbohydrates, lipids, or amino acids
At peak activity, energy is provided by anaerobic reactions
that generate lactic acid as a byproduct
10-6 Energy to Power Contractions
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Muscle Fatigue
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When muscles can no longer perform a required activity, they
are fatigued
Results of Muscle Fatigue
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Depletion of metabolic reserves
Damage to sarcolemma and sarcoplasmic reticulum
Low pH (lactic acid)
Muscle exhaustion and pain
10-6 Energy to Power Contractions
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The Recovery Period
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The time required after exertion for muscles to return to
normal
Oxygen becomes available
Mitochondrial activity resumes
© 2012 Pearson Education, Inc.
10-6 Energy to Power Contractions
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Lactic Acid Removal and Recycling
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The Cori Cycle
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The removal and recycling of lactic acid by the liver
Liver converts lactate to pyruvate
Glucose is released to recharge muscle glycogen reserves
10-6 Energy to Power Contractions
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The Oxygen Debt
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After exercise or other exertion:
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The body needs more oxygen than usual to normalize
metabolic activities
Resulting in heavy breathing
Also called excess postexercise oxygen consumption
(EPOC)
10-6 Energy to Power Contractions
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Heat Production and Loss
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Active muscles produce heat
Up to 70% of muscle energy can be lost as heat, raising body
temperature
© 2012 Pearson Education, Inc.
10-6 Energy to Power Contractions
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Hormones and Muscle Metabolism
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Growth hormone
Testosterone
Thyroid hormones
Epinephrine
10-7 Types of Muscles Fibers and Endurance
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Muscle Performance
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Force
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The maximum amount of tension produced
Endurance
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The amount of time an activity can be sustained
Force and endurance depend on
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The types of muscle fibers
Physical conditioning
10-7 Types of Muscles Fibers and Endurance
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Three Major Types of Skeletal Muscle Fibers
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Fast fibers
Slow fibers
Intermediate fibers
© 2012 Pearson Education, Inc.
10-7 Types of Muscles Fibers and Endurance
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Fast Fibers
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Contract very quickly
Have large diameter, large glycogen reserves, few
mitochondria
Have strong contractions, fatigue quickly
10-7 Types of Muscles Fibers and Endurance
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Slow Fibers
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Are slow to contract, slow to fatigue
Have small diameter, more mitochondria
Have high oxygen supply
Contain myoglobin (red pigment, binds oxygen)
10-7 Types of Muscles Fibers and Endurance
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Intermediate Fibers
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Are mid-sized
Have low myoglobin
Have more capillaries than fast fibers, slower to fatigue
10-7 Types of Muscles Fibers and Endurance
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Muscle Performance and the Distribution of Muscle Fibers
© 2012 Pearson Education, Inc.
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White muscles
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Mostly fast fibers
Pale (e.g., chicken breast)
Red muscles
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Mostly slow fibers
Dark (e.g., chicken legs)
Most human muscles
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Mixed fibers
Pink
10-7 Types of Muscles Fibers and Endurance
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Muscle Hypertrophy
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Muscle growth from heavy training
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Increases diameter of muscle fibers
Increases number of myofibrils
Increases mitochondria, glycogen reserves
Muscle Atrophy
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Lack of muscle activity
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Reduces muscle size, tone, and power
10-7 Types of Muscles Fibers and Endurance
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Physical Conditioning
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Improves both power and endurance
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Anaerobic activities (e.g., 50-meter dash, weightlifting)
© 2012 Pearson Education, Inc.
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Use fast fibers
Fatigue quickly with strenuous activity
Improved by:
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Frequent, brief, intensive workouts
Causes hypertrophy
10-7 Types of Muscles Fibers and Endurance
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Physical Conditioning
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Improves both power and endurance
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Aerobic activities (prolonged activity)
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Supported by mitochondria
Require oxygen and nutrients
Improves:
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Endurance by training fast fibers to be more like intermediate
fibers
Cardiovascular performance
10-7 Types of Muscles Fibers and Endurance
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Importance of Exercise
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What you don’t use, you lose
Muscle tone indicates base activity in motor units of skeletal
muscles
Muscles become flaccid when inactive for days or weeks
Muscle fibers break down proteins, become smaller and
weaker
© 2012 Pearson Education, Inc.
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With prolonged inactivity, fibrous tissue may replace muscle
fibers
10-8 Cardiac Muscle Tissue
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Cardiac Muscle Tissue
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Cardiac muscle cells are striated and found only in the heart
Striations are similar to that of skeletal muscle because the
internal arrangement of myofilaments is similar
10-8 Cardiac Muscle Tissue
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Structural Characteristics of Cardiac Muscle Tissue
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Unlike skeletal muscle, cardiac muscle cells (cardiocytes):
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Are small
Have a single nucleus
Have short, wide T tubules
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Have no triads
Have SR with no terminal cisternae
Are aerobic (high in myoglobin, mitochondria)
Have intercalated discs
10-8 Cardiac Muscle Tissue
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Intercalated Discs
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Are specialized contact points between cardiocytes
© 2012 Pearson Education, Inc.
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Join cell membranes of adjacent cardiocytes (gap junctions,
desmosomes)
Functions of intercalated discs:
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Maintain structure
Enhance molecular and electrical connections
Conduct action potentials
10-8 Cardiac Muscle Tissue
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Intercalated Discs
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Coordination of cardiocytes
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Because intercalated discs link heart cells mechanically,
chemically, and electrically, the heart functions like a single,
fused mass of cells
10-8 Cardiac Muscle Tissue
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Functional Characteristics of Cardiac Muscle Tissue
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Automaticity
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Contraction without neural stimulation
Controlled by pacemaker cells
Variable contraction tension
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Controlled by nervous system
Extended contraction time
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Ten times as long as skeletal muscle
Prevention of wave summation and tetanic contractions by cell
membranes
© 2012 Pearson Education, Inc.
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Long refractory period
10-9 Smooth Muscle Tissue
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Smooth Muscle in Body Systems
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Forms around other tissues
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In integumentary system
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Arrector pili muscles cause “goose bumps”
In blood vessels and airways
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Regulates blood pressure and airflow
In reproductive and glandular systems
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Produces movements
In digestive and urinary systems
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Forms sphincters
Produces contractions
10-9 Smooth Muscle Tissue
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Structural Characteristics of Smooth Muscle Tissue
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Nonstriated tissue
Different internal organization of actin and myosin
Different functional characteristics
10-9 Smooth Muscle Tissue
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Characteristics of Smooth Muscle Cells
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Long, slender, and spindle shaped
© 2012 Pearson Education, Inc.
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Have a single, central nucleus
Have no T tubules, myofibrils, or sarcomeres
Have no tendons or aponeuroses
Have scattered myosin fibers
Myosin fibers have more heads per thick filament
Have thin filaments attached to dense bodies
Dense bodies transmit contractions from cell to cell
10-9 Smooth Muscle Tissue
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Functional Characteristics of Smooth Muscle Tissue
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Excitation–contraction coupling
Length–tension relationships
Control of contractions
Smooth muscle tone
10-9 Smooth Muscle Tissue
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Excitation–Contraction Coupling
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Free Ca2+ in cytoplasm triggers contraction
Ca2+ binds with calmodulin
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In the sarcoplasm
Activates myosin light–chain kinase
Enzyme breaks down ATP, initiates contraction
© 2012 Pearson Education, Inc.
10-9 Smooth Muscle Tissue
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Length–Tension Relationships
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Thick and thin filaments are scattered
Resting length not related to tension development
Functions over a wide range of lengths (plasticity)
10-9 Smooth Muscle Tissue
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Control of Contractions
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Multiunit smooth muscle cells
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Connected to motor neurons
Visceral smooth muscle cells
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Not connected to motor neurons
Rhythmic cycles of activity controlled by pacesetter cells
10-9 Smooth Muscle Tissue
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Smooth Muscle Tone
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Maintains normal levels of activity
Modified by neural, hormonal, or chemical factors
© 2012 Pearson Education, Inc.
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