Vanders Human Physiology The Mechanisms of Body Function 13Th Edition Widmaier Solutions Manual Full Chapter PDF

Vanders Human Physiology The
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9: MUSCLE
This chapter is rewarding to teach because muscle physiology is relatively straightforward and because
one can present the material in sufficient depth to allow real understanding of it, even given the time
constraints of a general human physiology course. In addition, students have a natural interest in
learning about muscles and exercise physiology.
SECTION A: SKELETAL MUSCLE
9.1 Structure
Teaching/Learning Objectives: Following study of this section of the chapter, students should be able to:
Name the three types of muscle and indicate where they are found in the body.
Explain that a skeletal muscle is made up of many individual cells known as muscle fibers bound
together by connective tissue.
Recognize that skeletal muscles are usually attached to bones by bundles of collagen fibers called
tendons.
Describe the cytoplasm of each muscle fiber as containing cylindrical bundles of proteins called
myofibrils that are composed of the orderly overlapping of thick filaments, composed almost
entirely of the protein myosin, and thin filaments, principally composed of actin, troponin, and
tropomyosin, in a repeating pattern. Know that skeletal (and cardiac) muscle fibers are called
striated muscle because the overlapping of filaments results in a pattern of light and dark bands
or stripes.
Identify the various bands, lines, and zones within a sarcomere, stating which bands and zones
change in length during muscle shortening and which do not.
Note: Some students are puzzled by the fact that the I band, which spans two sarcomeres,
shortens, although emphasis is usually placed on the shortening of the sarcomere itself. One
source of confusion is the tendency to think of the thin filaments in adjoining sarcomeres as
being one long chain. They are not; they are connected to the Z line and not directly to each
other. As you can see from text Figure 95, during contraction both of the diagrammed
sarcomeres shorten simultaneously with the thin filaments in each being pulled toward their
respective M lines. This shortening is made possible by the anchorage of the thin filaments to
the Z lines. Likewise, although each sarcomere shortens, one end of the whole muscle remains
at a fixed point (the Zline on the right side of Figure 95) while the other end of the whole
muscle shortens towards it.
9.2 Molecular Mechanisms of Skeletal Muscle Contraction

Teaching/Learning Objectives: Following study of this section of the chapter, students should be able to
Define the term contraction, as the generation of tension in the muscle fiber and not necessarily
shortening.
Describe the sliding filament mechanism of contraction, including the chemical structure of the
thick and thin filaments and their relationship with each other. Recognize that actin has a
binding site for myosin, and that myosin heads, or cross‐bridges, have a binding site both for
actin and ATP. Recognize that the ATP binding site also serves as an enzyme that hydrolyzes
ATP.
Discuss the four steps in the crossbridge cycle, and describe the two separate functions
performed by ATP during the cycle, and appreciate that the same ATP molecule performs both
functions during a given cycle.
Discuss the roles of tropomyosin, troponin, and Ca2+ in regulating skeletal muscle contraction.
Describe the process of excitationcontraction coupling, including the role of the transverse
tubules, sarcoplasmic reticulum, dihydropyridine receptor and ryanodine receptor in the events
leading to Ca2+ release and the role of the Ca2+‐ATPase in contraction termination.
Define a motor unit, and recognize that whole muscles are composed of many motor units.
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Identify the functional anatomy of the neuromuscular junction and describe the events that
occur when an action potential reaches a motor neuron axon terminal. Compare and contrast
neuromuscular junctions and synapses.
Describe how the chemical agents curare, organophosphates, succinylcholine, rocuronium,
vecuronium, and botulinum toxin affect transmission at the neuromuscular junction.
9.3 Mechanics of SingleFiber Contraction

Differentiate among isometric contractions and the two types of isotonic contractions: concentric
contractions, and eccentric contractions.
Describe the properties of twitch contractions, including the latent period, contraction time, and
velocity of shortening.
Describe the load‐velocity relation of a muscle fiber.
Discuss how increasing the frequency of stimulation can lead to unfused and fused tetanic
contractions.
Discuss how the frequencytension relation is different for muscle fibers with different
contraction times.
Discuss why tetanic contractions produce greater tension than twitch contractions.
Describe the length‐tension relation.
9.4 Skeletal Muscle Energy Metabolism

State the three functions of ATP in skeletal muscle contraction and relaxation.
Discuss the three ways a muscle fiber can form ATP and describe the different fuel molecules
that are important for ATP production.
Describe how muscle fibers use ATP produced from creatine phosphate within the first few
seconds of increased activity.
Recognize that, at moderate levels of activity, most ATP used for muscle contraction is formed by
oxidative phosphorylation.
Discuss the contribution of glycolysis to ATP production during high‐intensity exercise.
Discuss the mechanisms thought to account for muscle fatigue during high‐intensity, short‐
duration exercise and during low‐intensity, long‐duration exercise.
Recognize the importance of fatigue as a mechanism for preventing the onset of rigor and muscle
damage.
9.5 Types of Skeletal Muscle Fibers

Describe the different characteristics of muscle fibers that allow them to be classified as fast
oxidativeglycolytic, slowoxidative, and fastglycolytic fibers.
Discuss how the differences in diameter of the various fiber types affect their strength of
contraction.
Differentiate among the fiber types with regard to their rate of fatigue.
9.6 WholeMuscle Contraction

Recognize that a whole muscle is made up of a combination of different types of muscle fibers
organized into motor units.
Recognize that the tension that can be developed in a whole muscle depends upon two factors:
the tension developed by each individual fiber and the number of active fibers.
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Describe how the number of active muscle fibers depends upon the number of fibers in a motor
unit and the recruitment of additional motor units.
Discuss why the order of recruitment of motor units depends on the diameter of the cell body of
the motor neuron.
Discuss why motor unit recruitment can also affect the shortening velocity of a whole muscle.
Describe the structural and functional effects of denervation atrophy, disuse atrophy, long‐
duration, low‐intensity exercise training, and short‐duration, high‐intensity exercise training.
Understand the extent to which different fiber types can change their properties.
Describe how muscles must be arranged around joints so that a limb can flex and extend.
Describe the advantages and disadvantages of the arrangement of the muscles, bones, and joints
in the body in lever systems.
9.7 Skeletal Muscle Disorders

Discuss the similarities and differences between the development of muscle cramps and
hypocalcemic tetany.
Describe the causes and symptoms of muscular dystrophy.
Describe the functional consequences and current treatments of myasthenia gravis.
Suggestion for class discussion: Worldclass athletes and muscle hypertrophy/changes in

function. An especially good example of the results of short‐duration, high‐intensity training on muscle
morphology is the Olympic speed skater, especially one who specializes in short‐distance races. Speed
skaters skate laps on an oval rink and always skate counter‐clockwise. Thus, they make only left turns
and the left leg must bear the brunt of the effort to turn. As a consequence of training, world‐class sprint
speed skaters often have a left leg that is larger in diameter than the right leg, sometimes by several
inches.
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LECTURE OUTLINE
I. Three types of muscle fibers (Fig. 9‐1)

II. Skeletal muscle
A. Structure (Fig. 9‐2)
1. Muscle, muscle fiber, myofibrils, myofilaments
2. Unit of contraction: the sarcomere (Fig. 9‐3)
B. Molecular mechanisms of contraction
1. The cross‐bridge cycle and the sliding filament mechanism (Fig. 9‐8)
2. Structure of thin filaments and regulation of contraction (Fig. 9‐9)
3. Excitation‐contraction coupling (Fig. 9‐10)
a. The roles of Ca2+ (Fig. 9‐12)
b. The neuromuscular junction leading to membrane excitation (Fig. 9‐15)
c. The motor unit and muscle contraction (Fig. 9‐13)
C. Mechanics of single‐fiber contraction
1. Isometric vs. isotonic contractions and tension vs. load
2 Properties of twitch contractions: latent period and contraction time (Fig. 9‐16)
3. Frequency‐tension relationship: summation and tetanus (Fig. 9‐20)
4. Load‐velocity relationship (Figs. 9‐17, 9‐18)
5. Length‐tension relationship (Fig. 9‐21)
D. Energy metabolism
1. Sources of ATP (Fig. 9‐22)
2. Muscle fatigue (Fig. 9‐23)
E. Types and properties of fibers (Table 9‐3)
F. Whole muscle contraction
1. Control of muscle tension and shortening velocity (Table 9‐4)
2. Muscle adaptation to exercise
3. Lever action of muscles and bones (Figs. 9‐27 through 9‐30)
G. Disorders
1. Muscle cramps vs. hypocalcemic tetany
2. Muscular dystrophy (Fig. 9‐31)
3. Myasthenia gravis
SECTION A REVIEW QUESTIONS
1. List the three types of muscle cells and their locations.

Skeletal, smooth, and cardiac. Most skeletal muscle is attached to bone, and its contraction moves and
supports the skeleton. Smooth muscle surrounds hollow organs and tubes, including the stomach, intestines,
urinary bladder, uterus, blood vessels, and lung airways. Cardiac muscle is the muscle of the heart.
2. Diagram the arrangement of thick and thin filaments in a striated muscle sarcomere, and label the
major bands that give rise to the striated pattern.
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A band
1/2 I band
Z line H zone
M line
Thin filament Thick filament
Sarcomere
3. Describe the organization of myosin, actin, tropomyosin, and troponin molecules in the thick and
thin filaments.
The myosin molecule is composed of two large polypeptide heavy chains and four smaller light chains.
These polypeptides combine to form a molecule that consists of two globular heads, containing heavy and
light chains, and a long tail formed by the two intertwined heavy chains. The tail of each myosin molecule
lies along the axis of the thick filament, and the two globular heads extend out to the sides, forming cross‐
bridges. Each globular head contains two binding sites, one for actin and one for ATP. The myosin molecules
in the two ends of each thick filament are oriented in opposite directions, such that their tail ends are
directed toward the center of the filament. The thick filament is primarily composed of myosin molecules.
The thin filament is principally composed of the protein actin, along with troponin and
tropomyosin. An actin molecule is a globular protein composed of a single polypeptide (a monomer) that
polymerizes with other actin monomers to form a polymer made up of two intertwined helical chains that
form the core of the thin filament. Each globular actin molecule contains a binding site for myosin.
Tropomyosin is a rodshaped molecule composed of two intertwined polypeptides with a length
approximately equal to seven actin monomers. Chains of tropomyosin are arranged end to end along the
actin thin filament. Troponin interacts with both tropomyosin and actin. It is composed of three subunits: I
(inhibitory), T (tropomyosinbinding), and C (Ca2+binding). One molecule of troponin binds to each
molecule of tropomyosin.
4. Describe the four steps of one cross‐bridge cycle.

A new crossbridge cycle begins with the binding of an energized myosin crossbridge (from step 4 of the
previous cycle) to actin. Binding of energized myosin to actin triggers the release of the strained
conformation of the energized bridge, which produces movement of the bound crossbridge (called the
power stroke) toward the center of the thick filament (step 2). During this movement, myosin releases the
ADP and Pi that had been bound to it in its energized state. At this point, myosin is firmly bound to actin.
Step 3 is the binding of a new molecule of ATP to the crossbridge, which causes myosin to decrease its
affinity for actin, and thus the crossbridge detaches. During step 4, the ATP bound to myosin is split by the
myosinATPase, thereby reforming the energized state of myosin and returning the crossbridge to its pre
power stroke position. The energized myosin with bound ADP and Pi is now free to bind to another molecule
of actin and start a new crossbridge cycle.
5. Describe the physical state of a muscle fiber in rigor mortis and the conditions that produce this
state.
Rigor mortis means “rigidity of death.” It is the gradual stiffening of skeletal muscles that begins several
hours after death and is caused by the depletion of ATP. In the absence of ATP, the breakage of the link
between actin and myosin does not occur because these events require ATP. The thick and thin filaments
remain bound to each other by immobilized crossbridges, producing a rigid condition in which the
filaments cannot be pulled past each other.
6. What three events in skeletal muscle contraction and relaxation depend on ATP?
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(1) Hydrolysis of ATP by myosin energizes the crossbridges, providing the energy for generation of
force.
(2) Binding of ATP to myosin dissociates crossbridges bound to actin, allowing the bridges to repeat
their cycle of activity. (Note that the same molecule of ATP performs both of these functions.)
(3) Hydrolysis of ATP by the Ca2+ATPase in the sarcoplasmic reticulum provides the energy for the
active transport of calcium ions into the lateral sacs of the reticulum, lowering cytosolic Ca2+ to prerelease
levels, which ends the contraction and allows the muscle fiber to relax.
7. What prevents cross‐bridges from attaching to sites on the thin filaments in a resting skeletal
muscle?
One molecule of troponin binds to each molecule of tropomyosin and regulates the access to myosin
binding sites on the seven actin monomers in contact with tropomyosin. In the resting skeletal muscle,
tropomyosin molecules partially cover the myosinbinding site on each actin monomer, thereby preventing
the crossbridges from making contact with actin. Each tropomyosin molecule is held in this blocking
position by a troponin molecule.
8. Describe the role and source of calcium ions in initiating contraction in skeletal muscle.
Troponin has a binding site for calcium ion. When Ca2+ is present in the cytosol, it binds to troponin and
causes a conformational change, which relaxes troponin's inhibitory grip and allows tropomyosin to move
away from the myosinbinding site on each actin molecule. Crossbridge cycling can continue to occur as
long as cytosolic Ca2+ concentration remains high enough. The source for cytosolic Ca2+ in skeletal muscle is
the sarcoplasmic reticulum.
9. Describe the location, structure, and function of the sarcoplasmic reticulum in skeletal muscle fibers.
The sarcoplasmic reticulum in muscle is homologous to the (smooth/agranular) endoplasmic reticulum
found in most cells. This structure forms a series of sleevelike segments around each myofibril. At the end of
each segment, there are two enlarged regions called lateral sacs that are connected to each other by smaller
tubular elements. The lateral sacs come in close contact with the Ttubules where the A bands and I bands of
the sarcomere meet. Ca2+ stored in the lateral sacs is released following membrane excitation.
10. Describe the structure and function of the transverse tubules.

Transverse tubules (Ttubules) are tubular structures separate from sarcoplasmic reticulum that
surround the myofibrils at the region of the sarcomere where the A band and I band meet. They are
continuous with the plasma membrane and invaginate deep into the muscle fiber passing between adjacent
lateral sacs. The lumen of each T tubule is continuous with the extracellular fluid. The membrane of the
tubule, like the plasma membrane, is able to propagate action potentials. Once initiated in the plasma
membrane, an action potential is rapidly conducted over the surface of the fiber and into its interior by way
of the Ttubules. An action potential in the Ttubule adjacent to the lateral sacs activates a voltagesensitive
protein in the Ttubule membrane that is physically linked to a Ca2+release channel in the membrane of the
lateral sacs. Depolarization of the Ttubule by an action potential, thus leads to the opening of the Ca2+
channels in the lateral sacs, allowing Ca2+ to diffuse from the Ca2+rich sarcoplasmic reticulum lumen into
the cytosol.
11. Describe the events that result in the relaxation of skeletal muscle fibers.
A contraction continues until Ca2+ is removed from troponin, and this is achieved by lowering the Ca2+
concentration in the cytosol back to its prerelease level. The membranes of the sarcoplasmic reticulum
contain primary activetransport proteins, Ca2+ATPases, that pump Ca2+ from the cytosol back into the
lumen of the reticulum. (This is the third function of ATP in skeletal muscle contraction and relaxation.) This
sequestering of calcium ion is slower than the process of Ca2+ release by diffusion, and so the cytosolic
concentration of Ca2+ remains elevated and the contraction continues for some time after a single action
potential.
12. Define a motor unit and describe its structure.
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A motor unit is a single motor neuron and all of the muscle fibers it innervates. Motor neurons are the
somatic efferent neurons of the peripheral nervous system. Their cell bodies are located either in the
brainstem or the spinal cord, and their axons are large in diameter and myelinated. Upon reaching a muscle,
the axon of a motor neuron divides into many branches, each branch forming a single junction with a
muscle fiber. A single motor neuron innervates many muscle fibers, but each muscle fiber is controlled by a
branch from only one motor neuron. Whole muscles are composed of many motor units.
13. Describe the sequence of events by which an action potential in a motor neuron produces an action
potential in the plasma membrane of a skeletal muscle fiber.
The junction of a motor neuron axon terminal with the muscle fiber it innervates is called a
neuromuscular junction, and the region of muscle fiber membrane that underlies the axon terminal has
special properties and is called the motor end plate. The axon terminals of motor neurons contain vesicles
(containing acetylcholine, ACh) similar to those found at synaptic junctions. When an action potential in the
motor neuron reaches the axon terminal, it depolarizes the membrane, opening voltagegated Ca2+ channels
and allowing calcium ions to diffuse into the axon terminal from the extracellular fluid. Calcium ions bind to
proteins that enable the membrane of the AChcontaining vesicles to fuse with the axon plasma membrane,
thereby releasing ACh into the synaptic cleft separating the axon terminal and the motor end plate.
ACh diffuses from the axon terminal to the motor end plate, where it binds to nicotinic acetylcholine
receptors. This binding produces a depolarizing potential in the motor end plate that is normally sufficient
to bring the muscle fiber plasma membrane adjacent to the end plate to its threshold potential, initiating an
action potential.
14. What is an end‐plate potential, and what ions produce it?

An end‐plate potential (EPP) is the local depolarization mentioned above that is generated in the
motor end plate by activation of nicotinic acetylcholine receptors, and is analogous to an excitatory
postsynaptic potential (EPSP). Like the EPSP, it is produced by the opening of ligandsensitive channels for
sodium and potassium ions. Because the electrochemical gradient for Na+ favors Na+ influx more than the
electrochemical gradient for K+ favors K+ efflux, the net result is a depolarization know as the EPP.
15. Compare and contrast the transmission of electrical activity at a neuromuscular junction with that at
a synapse.
As mentioned above, the endplate potential (EPP) is analagous to the EPSP at synapses. However, the
magnitude of the EPP is much greater than that of a single EPSP because the neuromuscular junction is
much larger in surface area than is a single synapse and neurotransmitter binds to many more receptors on
the motor end plate, opening many more ion channels. Thus, the effect of a neuromuscular junction is that of
an electrical synapse: An action potential in a motor neuron will normally generate action potentials in all
of the fibers it innervates. This is different from synaptic junctions between neurons, where multiple EPSPs
must occur in order for threshold to be reached and an action potential elicited in the postsynaptic
membrane.
Another difference between the neuromuscular junction and a synapse is that the motor end plate
receives information in the form of neurotransmitter from only one neuron. Thus, there is no convergence on
the motor end plate and no opportunity for integration of information, unlike the case with synapses. In
addition, the EPP is always excitatory; there is no change in motor endplate potential analogous to an IPSP.
Most neuromuscular junctions are located near the middle of each muscle fiber and newly generated action
potentials are propagated from this region in both directions toward the ends of the fiber. Finally, in
addition to receptors for ACh, the synaptic junction contains the enzyme acetylcholinesterase, which breaks
down ACh, just as it does at AChmediated synapses in the nervous system.
16. Describe isometric, concentric, and eccentric contractions.

Contraction refers to activation of the crossbridges, which thereby generate tension in a muscle.
Whether there is an accompanying change in muscle length depends upon the external forces acting on it.
The three types of contractions that can occur following activation of a muscle are: (1) isometric, in which
the muscle generates tension but does not change length; (2) isotonic concentric, in which the muscle
shortens and moves a load because the tension in the muscle exceeds the load; and (3) isotonic eccentric or
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lengthening, in which the external load on the muscle causes the muscle to lengthen during the period of
contractile activity because the load exceeds the tension produced by the muscle. These types of contractions
can occur in single fibers and in whole muscle.
17. What factors determine the duration of an isotonic twitch in skeletal muscle? An isometric twitch?
A twitch is the mechanical response of a single muscle fiber or a whole muscle to a single action
potential. The duration of an isotonic twitch is related to the magnitude of the load being moved: the
heavier the load, the shorter the duration of the twitch, partly due to the longer latent period with heavier
loads because more crossbridges need to bind in order to generate enough tension to lift a heavy load.
For an isometric twitch, duration depends upon the contraction time, which in turn partly depends
upon the Ca2+ATPase activity of the sarcoplasmic reticulum. This activity varies with fiber type: It is greater
(i.e., twitch duration is shorter) in fast fibers and less in slow fibers. Isometric twitch duration also depends
on how long it takes for crossbridges to complete their cycle and detach after removal of Ca2+ from the
cytosol.
18. What effect does increasing the frequency of action potentials in a skeletal muscle fiber have upon
the force of contraction? Explain the mechanism responsible for this effect.
Increasing the frequency of action potentials in a muscle fiber can increase the mechanical response
(tension), up to the level of maximum tetanic tension. This process is called summation. Summation is
possible because the duration of the action potential in skeletal muscle fibers is much shorter than the
duration of the twitch (12 ms vs. 100200 ms). Thus, a second action potential can be generated before the
tension has fallen to zero. When action potential frequency is very high, the tension becomes sustained at a
maximal level (fused tetanus).
The reason the relative tension is increased over that achieved by a single twitch is that, even
though a single action potential results in enough Ca2+ release to saturate troponin and make all of the
myosinbinding sites available on actin, time is required to bind the energized crossbridges to these sites.
Time is also required to transmit tension through the series of elastic structures, such as muscle tendons,
which must occur before the active force generated by the crossbridges can be applied to the load. For these
reasons, even though all of the binding sites are initially available during a single twitch, maximal tension
cannot be developed instantaneously. Calcium ions are pumped back into the sarcoplasmic reticulum almost
immediately after their release; therefore, many of the myosinbinding sites on actin have been covered
before crossbridges had a chance to bind.
In contrast, during a tetanic contraction, the successive action potentials each release Ca2+ from the
sarcoplasmic reticulum before all of the Ca2+ from the previous action potential has been pumped back into
the reticulum. This results in maintained high cytosolic Ca2+ concentrations and prevents a decline in the
number of available binding sites on the thin filaments. Since more crossbridges are able to bind as long as
Ca2+ is continually present, sustained tension, known as tetanus, can develop and the magnitude of the
tension depends on the frequency of action potentials sent to the muscle fiber.
19. Describe the length‐tension relationship in skeletal muscle fibers.
The amount of active tension that can be developed by a muscle fiber during contraction can be altered
by changing the length of the fiber before contraction. There is an optimal length, Lo, at which the muscle
fiber can generate maximal tension because at this length there is optimal overlap of thick and thin
filaments, allowing for maximal crossbridge cycling. When skeletal muscles are relaxed, the lengths of most
fibers are near Lo, and thus near the optimal lengths for force generation. If the fiber is stretched beyond this
length, some of the myosinbinding sites on the thin filaments will be pulled out of reach of the crossbridges,
and the maximal isometric tetanic tension that can be achieved will decline. Similarly, passive shortening of
the fiber will cause the actin filaments from opposite ends of the sarcomere to overlap and interfere with the
association of the myosin heads with the actin binding sites, leading to a decrease in the tension achieved.
Maximal active tension is achieved when the fiber is at its resting length, Lo. Fibers lose the ability
to generate active tension when stretched to about 170% of their resting length (although passive tension
at these lengths would be quite high) or shortened to 60% of it. Muscles in the body normally have a range
of movement from about 70–130% of their resting length due to their attachment to bones, meaning that
muscles are normally capable of generating at least half of the tension they can develop at Lo.
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20. Describe the effect of increasing load on a skeletal muscle fiber on the velocity of shortening.
The velocity at which a muscle fiber shortens decreases with increasing loads. The shortening velocity is
maximal when there is no load and zero when the load is equal to the maximal isometric tension. At loads
greater than the maximal isometric tension, the fiber will lengthen at a velocity that increases with load.
21. What is the function of creatine phosphate in skeletal muscle contraction?

When a muscle begins to contract, there is enough preformed ATP available to drive muscle activity for
only a few twitches. Energy is stored in muscle in a readily usable form called creatine phosphate (CP). ATP
can be generated rapidly by the phosphorylation of ADP by CP, a reaction that is catalyzed by creatine
kinase. The amount of ATP that can be formed from CP is limited to the amount of this highenergy
compound that is available, generally about five times as much CP as preformed ATP. Thus, for contractile
activity to be prolonged for more than a few seconds, other sources of ATP must be used.
22. What fuel molecules are metabolized to produce ATP during skeletal muscle activity?
Stored muscle glycogen is broken down to glucose, which is catabolized by glycolysis either
anaerobically or aerobically. As exercise proceeds, muscle fibers obtain glucose and fatty acids from blood.
The latter can be metabolized to produce ATP only in the presence of oxygen. Proteins can also be
catabolized into amino acids, which are then fed into the oxidative phosphorylation pathway for the
formation of ATP.
23. List the factors responsible for skeletal muscle fatigue.

Fatigue from highintensity, shortduration exercise involves at least three different mechanisms.
(1) Conduction failure: failure of the muscle action potential to be conducted into the fiber along the T
tubules, and thus a failure to release Ca2+ from the sarcoplasmic reticulum. The conduction failure results
from the buildup of potassium ions in the small volume of the Ttubule during the repolarization of each
successive action potential. This leads to a persistent depolarization of the membrane and eventually results
in failure of the membrane to produce action potentials due to inactivation of Na+ channels. (2) Lactic acid
buildup: elevated H+ concentration alters protein conformation and activity, including proteins such as
actin, myosin, and Ca2+ATPases. (3) Inhibition of crossbridge cycling: buildup of ADP and Pi within the
cytosol of muscle fibers may directly inhibit crossbridge cycling by the law of mass action.
With low intensity, longduration exercise, several processes have been implicated including the
three listed above, but at least two other mechanisms are thought to be more important: (1) Ryanodine
receptors become leaky during prolonged exercise, and the persistent elevation of cytosolic Ca2+ activates
proteases that degrade contractile proteins. (2) Although depletion of ATP is not a cause of fatigue, the
decrease other fuel substrates such as muscle glycogen correlates closely with fatigue onset.
24. What component of skeletal muscle fibers accounts for the differences in the fibers’ maximal
shortening velocities?
Fast and slow fibers contain different isoforms of the myosin ATPase that differ in the maximal rates at
which they split ATP. This difference, in turn, determines the maximal rate of crossbridge cycling and thus
the fiber’s maximal shortening velocity. Fibers containing myosin with high ATPase activity are classified as
fast fibers, and those containing myosin with lower ATPase activity are slow fibers.
25. Summarize the characteristics of the three types of skeletal muscle fibers.
In addition to having differences in myosin ATPase isoforms, skeletal muscle fibers may differ in the
major pathway they use to generate ATP—by oxidative phosphorylation or by glycolysis. Some fibers
contain numerous mitochodria and thus have a high capacity for oxidative phosphorylation. These fibers
are classified as oxidative fibers. Most of the ATP produced in these fibers is dependent upon blood flow to
deliver oxygen and fuel molecules to the muscle and so these fibers are surrounded by numerous small blood
vessels. They also contain large amounts of an oxygenbinding protein called myoglobin that increases the
rate of oxygen diffusion within the fiber and imparts a red color to the oxidative fibers; thus, oxidative fibers
are often called red muscle fibers. In contrast, glycolytic fibers have few mitochodria but possess a high
concentration of glycolytic enzymes and a large store of glycogen. Corresponding to their limited use of
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oxygen, these fibers are surrounded by relatively few blood vessels and contain little myoglobin. Thus, they
have a pale color and are often referred to as white muscle fibers.
On the basis of these two characteristics, three kinds of skeletal muscle fibers can be distinguished:
(1) Slow‐oxidative fibers (type I fibers) combine low myosinATPase activity with high oxidative
capacity.
(2) Fast‐oxidative‐glycolytic fibers (type IIa fibers) combine high myosinATPase activity with high
oxidative capacity and intermediate glycolytic capacity.
(3) Fast‐glycolytic fibers (type IIb fibers) combine high myosinATPase activity with high glycolytic
capacity.
The fibers differ from each other in other characteristics as well. Type IIb fibers have the largest
diameter, followed by type IIa fibers and then type I fibers. The number of thick and thin filaments per unit
of crosssectional area is about the same in all types of skeletalmuscle fibers. Accordingly,
type IIb fibers have a greater total number of thick and thin filaments than do the others and can thus
generate the greatest maximal tension, with type I fibers generating the least and the type IIa fibers being
intermediate.
The fibers also differ in their capacity to resist fatigue. Fastglycolytic fibers fatigue rapidly,
whereas slowoxidative fibers are very resistant to fatigue. Fastoxidativeglycolytic fibers have an
intermediate capacity to resist fatigue.
26. Upon what two factors does the amount of tension developed by a whole skeletal muscle depend?
The amount of tension developed by a whole skeletal muscle depends on: (1) the tension developed by
each individual fiber, which depends on the action potential frequency, fiber length, fiber diameter, and
fatigue; (2) the number of active fibers, which depends on the number of fibers per motor unit and the
number of active motor units.
27. Describe the process of motor‐unit recruitment in controlling (a) whole‐muscle tension and
(b) velocity of whole‐muscle shortening.
(a) Recruitment is the process of increasing the number of motor units that are active in a muscle at
any given time, and is achieved by increasing the excitatory input to the motor neurons. The greater the
number of active motor neurons, the more motor units will be recruited, and the greater the muscle tension.
(b) Similarly, the recruitment of motor units leads to a greater shortening velocity even in the face of
constant load because the load is shared among the active motor units. As more motor units become
recruited, the load for each individual unit will be effectively decreased and thus the velocity of whole
muscle shortening will increase.
28. During increases in the force of skeletal muscle contraction, what is the order of recruitment of the
different types of motor units?
Each motor unit contains only one fiber type, but whole muscles usually contain all three types of motor
units. The order of recruitment during skeletal muscle contraction is: motor units containing type I fibers
first; motor units containing type IIa fibers second; and motor units containing type IIb fibers last. The
reason for this order is that the diameter of the motor neuron cell body varies according to the diameter of
the muscle fibers it innervates. Motor neurons innervating type I fibers have the smallest diameter. Given the
same number of sodium ions entering a cell at a single excitatory synapse in a large diameter and a small
diameter motor neuron, the membrane of the small neuron will undergo a greater depolarization because
these ions will be distributed over a smaller surface area. Accordingly, given the same level of synaptic input,
the smallest motor neurons will be recruited first. The larger neurons will be recruited only as the level of
excitatory synaptic input increases. The next largest motor neurons, which innervate type IIa fibers, will be
activated to generate action potentials next, and finally, when synaptic input becomes strong enough, the
motor neurons innervating motor units containing type IIb fibers will be recruited.
29. What happens to skeletal muscle fibers when the motor neuron to the muscle is destroyed?
The regularity with which a muscle is used, as well as the duration and intensity of its activity, affect the
properties of the muscle. If the motor neurons to a muscle are destroyed, the denervated muscle fibers will
11
become progressively smaller in diameter, and the amount of contractile proteins they contain will decrease,
a condition known as denervation atrophy.
30. Describe the changes that occur in skeletal muscles following a period of (a) long‐duration, low‐
intensity exercise training; and (b) short‐duration, high‐intensity exercise training.
In contrast to the wasting seen when a muscle loses its nerve supply or is not used for an extended
period of time, increased contractile activity in the form of exercise has positive effects on muscle mass
and/or changes in their capacity for ATP production. Longduration, lowintensity exercise produces
increases in the number of mitochondria in the fibers that are recruited in this type of exercise (primarily
the oxidative fibers) and in the number of capillaries around them. These changes lead to an increase in the
capacity for endurance activity with a minimum of fatigue. Fiber diameter actually decreases slightly with
this type of training, accounting for the slender legs of marathon runners, for example.
Shortduration, highintensity activity affects primarily the fastglycolytic fibers, which are
recruited during strong contractions. With training, these fibers undergo an increase in diameter
(hypertrophy) due to an increased synthesis of actin and myosin filaments, which form more myofibrils. In
addition, their glycolytic activity is increased by increasing the synthesis of glycolytic enzymes. The result is
stronger, largerdiameter muscles, as seen in the legs of a sprinter or the legs and arms of a trained weight
lifter, however, the muscles have little capacity for endurance and fatigue rapidly.
Training of different types has little effect on the fast or slow myosinATPase characteristics of
muscle fibers, but it does change the biochemistry of the fibers, as described above. Thus, type IIb fibers can
be converted to type IIa fibers during endurance training and the opposite can occur during strength
training. If regular exercise ceases, the changes in the muscle that occurred as a result of the exercise will
slowly revert to their unexercised state.
31. How are skeletal muscles arranged around joints so that a limb can push or pull?
A contracting muscle exerts a force on bones through its connecting tendons. When the force is great
enough, the bone moves as the muscle shortens. A contracting muscle exerts only a pulling force, so that the
bones to which it is attached are pulled toward each other. In order for a limb to push as well as pull, there
must be at least two muscles on either side of a joint with their attachments to bones such that the
contraction of one will pull the joint in one direction and the contraction of the other will pull the joint in the
opposite direction. Flexion refers to the bending of a limb at a joint, and the muscles that cause this action
are called flexors. Extension refers to the straightening of the limb, and the muscles involved are called
extensors. Flexors and extensors are antagonists.
32. What are the advantages and disadvantages of the muscle‐bone‐joint lever system?
The disadvantage of the musclebonejoint lever system is a mechanical one and has to do with the
difference between the distance from the joint (e.g., the elbow) to the weight (e.g., in the hand) and the
distance from the joint to the point of insertion of the muscle (e.g., the biceps). In this example, the first
distance is 35 cm and the second is 5 cm. The product of the isometric tension generated by the muscle and
its distance from the joint must equal or exceed the product of the downward force of an object in the hand
and its distance away from the elbow in order to support or raise the weight in the hand. The 7fold
difference in the two distances described below means that the biceps must generate 70 kg of tension to
support a 10 kg weight.
However, the mechanical disadvantage that most muscle lever systems operate under is offset by
increased maneuverability. When the biceps shortens 1 cm, the hand moves through a distance of 7 cm.
Since the muscle shortens 1 cm in the same amount of time as the hand moves 7 cm, the velocity at which
the hand moves is 7 times greater than the rate of muscle shortening. Thus, the lever system amplifies the
velocity of muscle shortening.
12
SECTION B: SMOOTH AND CARDIAC MUSCLE
9.8 Structure of Smooth Muscle

Compare and contrast the structure of smooth muscle and skeletal muscle. Understand the
change in shape a smooth muscle fiber undergoes when it contracts.
9.9 Smooth Muscle Contraction and Its Control

Contrast the mechanisms by which cytosolic Ca2+ concentration regulates contractile activity in
smooth muscle and in skeletal muscle.
Contrast the sources for cytosolic Ca2+ in smooth and skeletal muscle, and identify the types of
inputs that can influence cytosolic Ca2+ concentration.
Describe a pacemaker potential.
Compare and contrast the neural control of skeletal muscle and smooth muscle.
Describe how neural and other types of controls of smooth muscle contractile activity can be
either stimulatory or inhibitory.
Contrast the properties of singleunit smooth muscle with those of multiunit smooth muscle, and
identify where in the body the different types of smooth muscle are found.
9.10 Cardiac Muscle

Compare and contrast cardiac muscle with skeletal muscle and smooth muscle.
Describe the function of intercalated disks and gap junctions.
Describe excitation‐contraction coupling in cardiac muscle including how L‐type Ca2+ channels
are involved.
Recognize that the strength of contraction can be varied by neurotransmitters or hormones.
Understand how the properties of L‐type Ca2+ channels contribute to the duration of a cardiac
twitch, and recognize why this relationship is important for normal cardiac function.
Describe how a single pacemaker potential causes the entire heart to beat.
LECTURE OUTLINE
I. Structure of smooth muscle
A. Smooth vs. skeletal muscle morphology
B. Arrangement of thick and thin filaments (Fig. 9‐33)
II. Excitation‐contraction coupling in smooth muscle
A. Regulation of contractile activity by Ca2+ (Fig. 9‐34)
B. Sources of cytosolic Ca2+
III. Regulation of smooth muscle contraction
A. Pacemaker potentials (Fig. 9‐36)
B. Control by the autonomic nervous system (Figs. 9‐37, 9‐38)
C. Control by hormones
D. Control by local factors
E. Control by stretch
IV. Types of smooth muscle
V. Cardiac muscle
A. Structure of cardiac muscle (Fig. 9‐39)
B. Excitation‐contraction coupling in cardiac muscle (Fig. 9‐40)
C. Timing of action potentials and twitch tension (Fig. 9‐41)
13
SECTION B REVIEW QUESTIONS
1. How does the organization of thick and thin filaments in smooth muscle fibers differ from that in
striated muscle fibers?
Smooth muscle, like skeletal muscle, has thick and thin filaments, but unlike the case with skeletal
muscle, they are not organized into myofibrils and there is no regular alignment of these filaments into
sarcomeres. Instead, the thin filaments are anchored either to the plasma membrane or to cytoplasmic
structures called dense bodies, and both sets of filaments are oriented slightly diagonally to the long axis of
the cell.
2. Compare the mechanisms by which a rise in cytosolic Ca2+ concentration initiates contractile activity
in skeletal, smooth and cardiac muscle fibers.
In skeletal muscle and cardiac muscle fibers, a rise in cytosolic Ca2+ concentration allows Ca2+ to bind to
troponin, which causes a conformational change in the molecule that relieves its inhibitory grip of
tropomyosin, and thus tropomyosin is able to move away from the crossbridge binding sites it was blocking
on the actin thin filament. In smooth muscle fibers, a rise in cytosolic Ca2+ concentration allows Ca2+ to bind
to calmodulin. The Ca2+calmodulin complex then binds to a protein kinase, myosin light‐chain kinase,
thereby activating the enzyme. The active protein kinase then uses ATP to phosphorylate myosin light
chains in the globular heads of myosin. This phosphorylation allows myosin crossbridges to bind to actin.
Hence, crossbridge activity in smooth muscle is turned on by Ca2+mediated changes in the thick filaments,
whereas in striated muscle (cardiac and skeletal), Ca2+ mediates changes in the thin filaments.
3. What are the two sources of Ca2+ that lead to the increase in cytosolic Ca2+ that triggers contraction
in smooth muscle?
Ca2+ that is stored in the sarcoplasmic reticulum and extracellular Ca2+ entering the cell through
plasmamembrane Ca2+ channels
4. What types of stimuli can trigger a rise in cytosolic Ca2+ in smooth muscle cells?
Action potentials in the plasma membrane of the fiber can be coupled with the release of Ca2+ ion from
the portions of the sarcoplasmic reticulum that are near the membrane. In addition, second messengers
released from the plasma membrane or generated in the cytosol in response to the binding of extracellular
chemical messengers to plasma membrane receptors can trigger the release of Ca2+ from more centrally
located sarcoplasmic reticulum.
There are voltage‐sensitive Ca2+ channels in the plasma membrane of smooth muscle cells that
respond to membrane depolarization to allow extracellular Ca2+ to diffuse into the cell. There are also Ca2+
channels in the membrane that are controlled by extracellular chemical messengers.
The types of inputs that affect cytosolic Ca2+ concentration in smooth muscle cells include:
(1) spontaneous electrical activity in the fiber’s plasma membrane;
(2) neurotransmitters released by autonomic neurons;
(3) hormones;
(4) locally produced changes in the chemical composition of the extracellular fluid surrounding the
fiber, and
(5) stretch.
5. What effect does a pacemaker potential have on a smooth muscle cell?

Some types of smooth muscle fibers do not maintain a constant resting potential. Instead, they
gradually depolarize to threshold potential whereupon they produce an action potential—in the absence of
any neural or hormonal input. The potential change during the spontaneous depolarization to threshold is
called the pacemaker potential. Following repolarization, the membrane again begins to depolarize, so that
a sequence of action potentials occurs, producing a tonic state of contractile activity.
6. In what ways does the neural control of smooth muscle activity differ from that of skeletal muscle?
14
Each skeletal muscle fiber is controlled by a single motor neuron that releases the excitatory
transmitter acetylcholine at its neuromuscular junction with the motor end plate.
Smooth muscle fibers do not have specialized endplate regions. They receive neurotransmitters by
diffusion from the varicosities of the branches of postganglionic autonomic neurons that enter the region of
smooth muscle fibers. Varicosities from a single axon may be located along several muscle fibers, and a
single muscle fiber may be located near varicosities belonging to parasympathetic and sympathetic
postganglionic neurons. Therefore, a single smooth muscle fiber may be influenced by neurotransmitter
from more than one neuron.
Also unlike the case with neural control of skeletal muscle, smooth muscle fibers may be stimulated
to contract by some neurotransmitters and be inhibited from contracting by others.
7. Describe how a stimulus may lead to the contraction of a smooth muscle cell without a change in the
plasma membrane potential.
Smooth muscle cells respond to extracellular chemical messengers in a variety of ways. Some of these
messengers elicit responses in the plasma membrane that generate second messengers, such as inositol
triphosphate, which cause Ca2+ to be released from the sarcoplasmic reticulum. Thus, contraction can be
initiated without a change in membrane potential.
8. Describe the differences between single‐unit and multiunit smooth muscle.

As the name suggests, single‐unit smooth muscle cells respond to stimulation as if they were a single
unit—i.e., their electrical and mechanical activities are synchronized. This occurs because each muscle fiber
is linked to adjacent fibers by gap junctions, through which action potentials occurring in one cell are
propagated to other cells by local currents. Some of the fibers in a singleunit muscle are pacemaker cells
that spontaneously generate action potentials that are propagated by way of gap junctions into adjacent
cells that do not spontaneously generate them (the majority of the cells).
The contractile activity of singleunit smooth muscles can be altered by nerves, hormones, local factors,
and stretch. The extent of innervation of these muscles varies in different organs, but generally the nerve
terminals are restricted to the regions of the muscle that contain the pacemaker cells. By regulating the
frequency of the pacemaker cells’ action potentials, the activity of the entire muscle can be controlled.
Multiunit smooth muscles have no or few gap junctions between fibers, each fiber responds
independently of its neighbors, and the muscle behaves as multiple units. Multiunit smooth muscles are
richly innervated by branches of the autonomic nervous system. The contractile response of the whole
muscle depends on the number of muscle fibers that are activated and on the frequency of nerve stimulation.
Although stimulation of some of the nerve fibers to the muscle can lead to depolarization and a contractile
response, action potentials do not occur in most multiunit smooth muscles. These muscles also respond to
hormones by increasing or decreasing contractile activity, but they do not respond to stretch.
9. Compare and contrast the physiology of cardiac muscle with that of skeletal and smooth muscles.
Cardiac muscle often combines properties of both skeletal and smooth muscle. Similar to smooth muscle, the
source of Ca2+ in cardiac muscle contractions is from both the sarcoplasmic reticulum and extracellular
sources. Both skeletal and cardiac muscles contain DHP receptors. In skeletal muscles, DHP receptors are
voltagesensors that tug open the ryanodine receptors embedded within the sarcoplasmic reticulum
membrane in response to propagating action potentials. In cardiac muscles, DHP receptors are also called
Ltype Ca2+ channels. Here they function as channels, allowing Ca2+ to enter the cytosol from the ECF. This
"trigger Ca2+" binds to ryanodine receptors and causes these channels to open. In addition, unlike smooth
muscle, both cardiac and skeletal muscle contain transverse tubules. In skeletal and cardiac muscle Ca2+
binds troponin to regulate contraction while in smooth muscle Ca2+ binds to calmodulin, and the complex
activates myosin light chain kinase, which phosphorylates the myosin crossbridge head allowing for
contraction to take place. Neurotransmitters can control all three types of muscles; however, in smooth and
cardiac muscle, innervation can be either stimulatory or inhibitory while it is only stimulatory in skeletal
muscle. In addition, hormones can control the contraction of smooth and cardiac muscles but not skeletal
muscle. Gap junctions are present in singleunit smooth muscle and cardiac muscle, and they permit the
passage of action potentials from cell to cell and thus allow for coordinated contractions. Also, both single
unit smooth and cardiac muscles contain pacemaker cells which spontaneously generate action potentials.
15
Finally, cardiac muscles are prevented from having tetany while skeletal muscle and smooth muscle can
undergo tetany.
10. Explain why cardiac muscle cannot undergo tetanic contractions.

Cardiac muscle cells contain voltagegated Ca2+ channels, called Ltype Ca2+ channels. These channels
remain open for long periods of time upon stimulation, allowing a continuous flow of Ca2+ into the cell. In
fact, the action potential duration of a cardiac muscle cell lasts as long as the twitch tension (200300 ms).
While Ltype Ca2+ channels remain open additional stimuli cannot have an effect on the cardiac cell, thus
cardiac cells cannot undergo tetany. This is necessary for the normal contraction and relaxation of the
heart and hence pumping of blood.
ANSWERS TO TEST QUESTIONS
91 a A single skeletal muscle fiber, or cell, is composed of many myofibrils.
92 e The dark stripe in a striated muscle that constitutes the A band results from the aligned thick
filaments within myofibrils, so thick filament length is equal to A‐band width.
93 b As filaments slide during a shortening contraction, the I band becomes narrower, so the distance
between the Z line and the thick filaments (at the end of the A band) must decrease.
94 d DHP receptors act as voltage sensors in the T‐tubule membrane and are physically linked to
ryanodine receptors in the sarcoplasmic reticulum membrane. When an action potential depolarizes the
T‐tubule membrane, DHP receptors change conformation and trigger the opening of the ryanodine
receptors. This allows Ca2 to diffuse from the interior of the sarcoplasmic reticulum into the cytosol.
95 c In an isometric twitch, tension begins to rise as soon as excitation–contraction is complete and
the first cross‐bridges begin to attach. In an isotonic twitch, excitation–contraction coupling takes the
same amount of time, but the fiber is delayed from shortening until after enough cross‐bridges have
attached to move the load.
96 b In the first few seconds of exercise, mass action favors transfer of the high‐energy phosphate
from creatine phosphate to ADP by the enzyme creatine kinase.
97 d Fast‐oxidative‐glycolytic fibers are an intermediate type that are designed to contract rapidly
but to resist fatigue. They utilize both aerobic and anaerobic energy systems; thus, they are red fibers
with high myoglobin (which facilitates production of ATP by oxidative phosphorylation), but they also
have a moderate ability to generate ATP through glycolytic pathways. (Refer to Table 9–3.)
98 c In smooth muscle cells, dense bodies serve the same functional role as Z lines do in striated
muscle cells—they serve as the anchoring point for the thin filaments.
99 b When myosin‐light‐chain kinase transfers a phosphate group from ATP to the myosin light
chains of the cross‐bridges, binding and cycling of cross‐bridges is activated.
910 d Stretching a sheet of single‐unit smooth muscle cells opens mechanically gated ion channels,
which causes a depolarization that propagates through gap junctions, followed by Ca2 entry and
contraction. This does not occur in multiunit smooth muscle.
911 e The amount of Ca2 released during a typical resting heart beat exposes less than half of the thin
filament cross‐bridge binding sites. Autonomic neurotransmitters and hormones can increase or
decrease the amount of Ca2 released to the cytosol during EC coupling.
16
ANSWERS TO QUANTITATIVE AND THOUGHT QUESTIONS
91 Under resting conditions, the myosin has already bound and hydrolyzed a molecule of ATP, resulting
in an energized molecule of myosin (M · ADP · Pi). Because ATP is necessary to detach the myosin cross‐
bridge from actin at the end of cross‐bridge movement, the absence of ATP will result in rigor mortis, in
which case the cross‐bridges become bound to actin but do not detach, leaving myosin bound to actin (A 
M).
92 The distance from the elbow joint to the hand (30 cm) is 10 times greater than the distance from the
elbow joint to the biceps insertion (3 cm), so her hand will move 20 m/sec when her biceps contracts at
2 m/sec (see Figure 9.30).
93 The length–tension relationship states that the maximum tension developed by a muscle decreases
at lengths below L0. During normal shortening, as the sarcomere length becomes shorter than the
optimal length, the maximum tension that can be generated decreases. With a light load, the muscle will
continue to shorten until its maximal tension just equals the load. No further shortening is possible
because at shorter sarcomere lengths the tension would be less than the load. The heavier the load, the
less the distance shortened before reaching the isometric state.
94 Maximum tension is produced when the fiber is (a) stimulated by an action potential frequency that
is high enough to produce a maximal tetanic tension and (b) at its optimum length L0, where the thick
and thin filaments have overlap sufficient to provide the greatest number of cross‐bridges for tension
production.
95 Moderate tension—for example, 50 percent of maximal tension—is accomplished by recruiting

sufficient numbers of motor units to produce this degree of tension. If activity is maintained at this level
for prolonged periods, some of the active fibers will begin to fatigue and their contribution to the total
tension will decrease. The same level of total tension can be maintained, however, by recruiting new
motor units as some of the original ones fatigue. At this point, for example, one might have 50 percent of
the fibers active, 25 percent fatigued, and 25 percent still unrecruited. Eventually, when all the fibers
have fatigued and there are no additional motor units to recruit, the whole muscle will fatigue.
96 The oxidative motor units, both fast and slow, will be affected first by a decrease in blood flow
because they depend on blood flow to provide both the fuel—glucose and fatty acids—and the oxygen
required to metabolize the fuel. The fast‐glycolytic motor units will be affected more slowly because they
rely predominantly on internal stores of glycogen, which is anaerobically metabolized by glycolysis.
97 Two factors lead to the recovery of muscle force. (a) Some new fibers can be formed by the fusion
and development of undifferentiated satellite cells. This will replace some, but not all, of the fibers that
were damaged. (b) Some of the restored force results from hypertrophy of the surviving fibers. Because
of the loss of fibers in the accident, the remaining fibers must produce more force to move a given load.
The remaining fibers undergo increased synthesis of actin and myosin, resulting in increases in fiber
diameter and consequently their force of contraction.
98 In the absence of extracellular calcium ions, skeletal muscle contracts normally in response to an
action potential generated in its plasma membrane because the Ca2 required to trigger contraction
comes entirely from the sarcoplasmic reticulum within the muscle fibers. If the motor neuron to the
muscle is stimulated in a calcium‐free medium, however, the muscle will not contract because the influx
of Ca2 from the extracellular fluid into the motor nerve terminal is necessary to trigger the release of
acetylcholine that in turn triggers an action potential in the muscle.
In a calcium‐free solution, smooth muscles would not respond either to stimulation of the nerve or to
the plasma membrane. Stimulating the nerve would have no effect because Ca2 entry into presynaptic
17
terminals is necessary for neurotransmitter release. Stimulating the smooth muscle cell membrane
would also not cause a response in the absence of Ca2 because in all of the various types of smooth
muscle, Ca2 must enter from outside the cell to trigger contraction. In some cases, the external Ca2
directly initiates contraction, and in others it triggers the release of Ca2 from the sarcoplasmic reticulum
(calcium‐induced Ca2 release).
99 The simplest model to explain the experimental observations is as follows. Upon parasympathetic
nerve stimulation, a neurotransmitter is released that binds to receptors on the membranes of smooth
muscle cells and triggers contraction. The substance released, however, is not acetylcholine (ACh) for the
following reason.
Action potentials in the parasympathetic nerves are essential for initiating nerve‐induced
contraction. When the nerves were prevented from generating action potentials by blockage of their
voltage‐gated Na channels, there was no response to nerve stimulation. ACh is the neurotransmitter
released from most, but not all, parasympathetic endings.
When the muscarinic receptors for ACh were blocked, however, stimulation of the parasympathetic
nerves still produced a contraction, providing evidence that some substance other than ACh is being
released by the neurons and producing contraction.
910 Elevation of extracellular fluid Ca2 concentration would increase the amount of Ca2 entering
the cytosol through L‐type Ca2 channels. This would result in a greater depolarization of cardiac muscle
cell membranes during action potentials. The strength of cardiac muscle contractions would also be
increased because this larger Ca2 entry would trigger more Ca2 release through ryanodine receptor
channels, and consequently there would be a greater activation of cross‐bridge cycling.
9.11 In order for unfused tetanus to occur, action potentials must occur more closely in time than the
duration of a twitch cycle. Frequency is the inverse of cycle duration, so to produce unfused tetanus,
action potentials must occur at a frequency greater than 1/0.040 seconds, or 25 action potentials per
second.
ANSWERS TO GENERAL PRINCIPLES ASSESSMENTS
9.1 The control of cardiac muscle pacemaker cell activity by sympathetic and parasympathetic
neurotransmitters is an excellent example of the principle that most physiological functions are controlled
by multiple regulatory systems, often working in opposition.
9.2 The forward motion of cross‐bridges during the cross‐bridge cycle (power stroke) is associated with
a chemical reaction in which ADP and Pi are released as products (see Step 2 in Figure 9.15). During
high‐frequency stimulation of muscles when cross‐bridges cycle repeatedly, the concentrations of ADP
and Pi build up in the muscle cytosol. Due to the law of mass action, the buildup of these products
inhibits the rate of the chemical reaction, and thus the powerstroke of the cross‐bridge cycle. This
contributes to the reduction of contraction speed and force that occurs when muscles are fatigued.
9.3 The principle of controlled exchange of materials between compartments and across membranes is
demonstrated by the movements of Ca2+ and other ions involved in the skeletal muscle EC‐coupling
mechanism (see Figure 9.40). Controlled movement of Na+, K+ and Ca2+ across muscle cell plasma
membranes maintains the resting membrane potential and allows the generation and propagation of
action potentials. Sequestering Ca2+ in the sarcoplasmic reticulum allows the resting state of muscle to
be maintained, until controlled release of Ca2+ into the cytosol activates cross‐bridge cycling and muscle
contraction. The termination of muscle contraction requires the return of Ca2+ into the sarcoplasmic
reticulum and extracellular fluid.
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calling out, “Abdullah, eat; for you are a hungry Kafir.” I found that,
during my absence, only one kafila had arrived from Bornou,—the
same which had brought me the letters, along with three bottles of
port wine, and some gunpowder, from Major Denham. Hat Salah,
among other news, mentioned that old Jacob, my servant, had been
in great distress for my safety during my absence; and that a female
slave of El Wordee’s, who was much attached to him, had lost her
reason on hearing we were gone to Youri, and in this unhappy state,
having thrown herself into a well, she had broken one of her arms.
May 23.—Cool and cloudy. I was visited by all the principal Arabs
who were in Kano; amongst the rest old Hadje Boo Zaied, who has
ever been our stanch friend, and was a very worthy man. He
begged, with great earnestness, that I would not acquaint the sheikh
of Bornou or the bashaw of Tripoli of Bello’s behaviour to Hadje Ali at
Sackatoo. For Boo Zaied’s sake, I promised to screen him, unless
questions were expressly put to me concerning his conduct, when I
must speak the truth; for he had behaved to me both like a fool and a
knave.
May 25.—To-day I paid up my servants’ wages, at the rate of four
dollars a month, but reduced them in future one half; notwithstanding
which, they were all glad to remain in my service.
May 26.—I waited on the governor, who received me with marked
kindness, and inquired particularly after the health of the sultan and
of the gadado, and how I had fared in crossing the Gondamee, the
river between Futche and Sackatoo.
May 30.—Clear and sultry. I was earnestly solicited by the people
to refer to my books, and to ascertain if the new moon would be
seen to-day; which much longed-for event, I assured them, would
take place after sunset, if the evening was clear. This anxiety was
occasioned by the fast of the Rhamadan, then terminating, and the
Aid, or great feast, immediately commencing. The evening turning
out cloudy, all were in low spirits; but at midnight a horseman arrived
express to acquaint the governor that the new moon had been
visible.
May 31.—After the arrival of the horseman, nothing was heard but
the firing of musketry and shouts of rejoicing.—Paying and receiving
visits now became a serious occupation. In the morning,
accompanied by Hat Salah, I went on horseback to pay my respects
to the governor. I accepted his invitation to ride out with him,
according to their annual custom; and we proceeded to an open
space within the city walls, amid skirmishing and firing of muskets,
attended by his people on horseback, and the Arabs and principal
townsfolk dressed in their gayest raiments,—all who could possibly
muster a horse for the occasion being mounted. The most
conspicuous person in the whole procession was a man on
horseback in quilted armour, who rode before the governor bearing a
two-handed sword. On reaching the plain, the governor made a
speech to the people, declaring his intention to attack Duntungua,
when he expected every man to exert his utmost prowess. Their
sons too should not, as in times past, be left behind, but would
accompany them to the war, and learn to fight the battles of their
country under the eyes of their parents. Afterwards we rode home in
the same order. All work was laid aside for three days. Men, women,
and children, in their finest clothes, paraded through the town; a
number of slaves were also set free, according to the custom of
Mahometans at this holy season. The owner of my house freed
fifteen.
June 1.—I visited the governor, to take leave. He was very kind,
and after inquiring if I should ever return, begged me to remember
him to his friend the sheikh El Kanemy, and expressed his hope I
would give a favourable account of the people I had visited. I
assured him, as to the last particular, I could not do otherwise, as I
had every where experienced the greatest civility. He then repeated
the Fatha, and I bade him farewell.
June 3.—At ten in the morning I left Kano, and was accompanied
some miles by Hadje Hat Salah and all my friends on horseback.
Before Hat Salah left me, he called all my servants before him, and
told them he trusted they would behave well and faithfully; for, as
they had seen, I was the servant of a great king, the friend of the
bashaw of Tripoli, and had been passed from one sultan to another;
consequently any misbehaviour of theirs, on a complaint from me,
would be severely punished. We only travelled a short way before
halting, for the heat of the day, under a shady tree. In the afternoon
we again set forward, and at sunset encamped outside the town of
Duakee.
June 4.—This morning we passed through the walled town of
Sockwa, which is now reduced to a few huts inhabited by slaves;
and halting for the heat of the day under a tamarind tree, we pitched
our tents at sunset under the walls of Girkwa, not far from the banks
of the river. The people were dancing in honour of the Aid. The
dance was performed by men armed with sticks, who springing
alternately from one foot to the other, while dancing round in a ring,
frequently flourished their sticks in the air, or clashed them together
with a loud noise. Sometimes a dancer jumped out of the circle, and
spinning round on his heel for several minutes, made his stick whirl
above his head at the same time with equal rapidity; he would then
rejoin the dance. In the centre of the ring there were two drummers,
the drums standing on the ground. They were made of a hollow
block of wood about three feet high, with a skin drawn tensely over
the top by means of braces. A great concourse of natives were
assembled to witness the exhibition.
June 5.—Morning cloudy. At six in the morning we left Girkwa,
and reposing ourselves during the heat of the day under some
tamarind trees among the villages of Nansarina, we encamped at
sunset in the woods. The inhabitants were now very busy in the
fields planting grain. Their mode of planting it is very simple. A man
with a hoe scrapes up a little mould at regular intervals, and is
followed by a woman carrying the seed, of which she throws a few
grains into each hole, and treads down the mould over them with her
feet.
June 6.—At noon we halted in the town of Sangeia, the governor
of which was at Kano; so I fortunately escaped the pain of hearing
his squeaking voice. We encamped for the night in the woods.
June 7.—At one in the afternoon we halted outside the town of
Katungwa. At sunset two horsemen arrived at full gallop, with the
news of the governor of Kano having taken a town, at a very short
distance to the north, from the rebel Duntungua.
June 8.—Every where the inhabitants were busily employed
clearing the ground, and burning the weeds and stubble, preparatory
to sowing grain. We sheltered ourselves from the mid-day heat
under the shade of a tamarind tree, in the province of Sherra, and
halted for the night outside the town of Boosuea. A son of the
governor of Sherra was here, attended by a number of horsemen,
and a band of music. He drank coffee with me, and I was in turn
regaled with music the greater part of the night. The instruments
were chiefly flutes and long wooden pipes, called by the natives
frum-frum.
June 9.—At sunset we arrived at the town of Dugwa.
June 10.—At daybreak we left Dugwa, and travelled through a
thickly wooded country. It rained all day, and we also had some
thunder and lightning. At seven in the evening we arrived at Murmur.
I heard, at Kano, that a kafila of Arabs, belonging to Augela, had
destroyed the clay wall around Dr. Oudney’s grave, and made a fire
over it, telling the inhabitants he was a Kafir. This report, to my great
regret, I found to be true.
June 11.—At sunrise I sent for the governor, to inquire who had
committed the outrage, when he protested it was the Arabs, and not
the people of the town. I felt so indignant at this wanton act of
barbarity, I could not refrain from applying my horsewhip across the
governor’s shoulders, and threatened to report him to his superior,
the governor of Katagum, and also to despatch a letter on the
subject to the sultan, unless the wall was immediately rebuilt: which,
with slavish submission, he promised faithfully to see done without
delay. During my halt at noon, near Katagum, I sent Dumbojee
forward to inform Duncawa, the governor, of my return. In the
afternoon I heard that he was on his way to meet me; and I had
scarcely left my resting-place before he made his appearance,
attended by about thirty horsemen, who, when they saw me, came
up at full gallop, brandishing their spears. I presented the governor
with a hundred Goora nuts, every one of which he distributed
amongst his people. He gave me many very hearty welcomes, and
made numerous inquiries about Bello, and his behaviour to me. He
and his people now galloped into the town, yelling and skirmishing;
and although the governor had been sick for some time past, he
appeared as lively and cheerful as any of them. On entering
Katagum I was lodged in my old quarters, and was immediately
visited by my old friend Hameda, the Tripoline merchant, who was
still here. I invited him to accompany me to Tripoli, as the late Dr.
Oudney had advised him; but he excused himself, on the plea of
being unable to collect his outstanding debts from his numerous
creditors, who were scattered all over the country.
June 12.—Warm and sultry. Duncawa remained with me all day,
and informed me, that he had the sultan’s orders to conduct me to
Kouka, in Bornou. This mark of respect I positively declined, both on
account of his recent illness, and also lest his presence might give
umbrage to the sheikh; but agreed to accept from him an escort
through the Bede territory. I assured him, when once in Bornou, that
I felt myself as safe as in his house. If he insisted, however, on
somebody accompanying me, he might, if he pleased, send one of
his principal people. I made a formal complaint of the insult
committed to Dr. Oudney’s grave,—enforcing, in the strongest terms,
the disgrace of disturbing the ashes of the dead, whose immortal
part was now beyond the power of malignant man. He frankly
acknowledged the enormity of the act, and faithfully promised to
have the wall rebuilt,—even offering to send for the governor of
Murmur, and have him punished; but, at the same time, begging me
not to acquaint the sultan of the occurrence. I expressed my reliance
on his assurances, but apprised him I must inform the gadado of the
affair. I afterwards spent the evening with Hameda.
June 13.—There was a fresh breeze in the morning; but it
afterwards began to rain. Duncawa being laid up from lameness, I
had a day’s rest, and again spent the evening with Hameda. The
conversation turning on the trustworthiness of slaves, he mentioned
to me, that his servants never knew in what apartment of his house
he slept; and that he even lay with a dagger, and loaded pistols,
under his pillow, lest he should be murdered by his female slaves.
He also acquainted me, that almost all the Arabs did the same; for it
was chiefly females whom they had reason to fear, the master being
often strangled at night by the women of his household.
June 14.—Duncawa visited me again, and made me a present of
two tobes, two sheep, and a large quantity of Guinea corn, and gave
a tobe to each of my servants. I presented him with six hundred
Goora nuts, having brought a large supply of them from Kano.
June 15.—I had every thing prepared for continuing my journey,
but Duncawa pressed me to spend another day with him, and I
availed myself of the delay to write to Bello and the gadado. I
returned my humble thanks to the former for his protection and
favour while I sojourned in his territories; and, in acknowledging the
uniform kindness of the latter, I did not fail to acquaint him of the
outrage committed on Dr. Oudney’s grave. I delivered these letters to
the charge of Dumbojee, who, having fulfilled his orders, took leave
of me here, having first made him a present of a couple of tobes and
forty dollars. My guide, Mahomed Dumbojee, had now become rich
and gay, having a numerous train of attendants; for at every town
where we halted, the governor was bound in courtesy to make him a
present, in token of respect for the sultan.
Having sent my camels forward, I went to bid farewell to
Duncawa, who was still confined to his house by illness. He made
me breakfast with him. Our breakfast consisted of a sheep’s head,
singed in the same manner as is practised in Scotland—a sheep’s
fry—and bread and milk. I was accompanied across the Yeou by my
friend Hameda, and Duncawa’s horsemen, who all wished to be
allowed to attend me to Sansan; but I excused myself from this
guard of honour, at once troublesome and expensive, by pretending
it was unlucky to go beyond the banks of a river with a friend.
Attended only by one of Duncawa’s principal men, I passed the thick
woods on the bank of the river, and, halting under a tamarind tree
during the heat of the day, I encamped towards evening at a village
called Mica. The inhabitants were all very busy in the fields sowing
gussub. They brought me, however, an abundant supply of milk, and
repeated inquiries were made after Bello’s health; for although they
recently belonged to Bornou, of which country they are natives, they
entertain, nevertheless, a great respect for their new sultan.
June 17.—I started at daylight, and, as the weather was cloudy
and rather windy, I did not halt before reaching Sansan. I was here
provided with very indifferent accommodation; but, on threatening I
would encamp outside the town, the governor received me into his
own house, according to Duncawa’s orders, and also made me a
present of a sheep. At night there was a violent storm, with thunder
and lightning. The poor lad Joseph, who had been hired at Kouka by
the late Dr. Oudney to tend the camels, was out all night with them.
Being a native of Fezzan, and half an idiot, he was here considered
a holy man, and I still retained him in my service out of charity. It was
he who gave me an account of the people of Bede, as he had been
a slave among them; and related his story with such artless
simplicity, that I implicitly rely on its correctness.
June 18.—Cool and cloudy. I heard to-day of a courier being
delayed on his route, by his camel’s being knocked up; and as
Duncawa was also preparing a present for the sheikh El Kanemy, I
postponed my departure yet another day.
June 19.—At eleven in the forenoon the courier arrived, bringing a
sabre as a present for the sultan Bello, and letters from Major
Denham, the consul at Tripoli, and the secretary of state. Accordingly
at mid-day I set off on my return to Katagum, in order to have the
sword forwarded to Bello by Duncawa.
At ten in the morning I entered Katagum, and immediately waited
on Duncawa to acquaint him with the cause of my return. I showed
him the sword, and explaining to him the manner of attaching the
belt, he expressed himself in terms of the highest admiration of both
sheath and sabre; and looking again and again at the ornaments, he
frequently asked, “Is not this all gold?” He sent instantly for the cadi,
who wrote a letter in my name to Bello, and a courier was
despatched with it and the sword. In the evening, another thunder-
storm, with much rain.
June 21.—At one in the afternoon I arrived again at Sansan.
June 22.—Clear and sultry. I was further detained on account of
the present for the sheikh not being ready.
June 23.—Morning cloudy. At seven in the morning I left Sansan,
attended by part of the escort which was to conduct me through the
Bede territory, and was obliged to stop about noon at the village of
Girkwa, by a violent attack of ague and bilious vomiting. Previous to
starting, I was joined by two merchants of Tripoli, who had been at
Kano, and begged to be allowed to place themselves under my
protection during this perilous part of the journey. In the afternoon
Hadje Fudor, the governor of Sansan, arrived with the remainder of
the escort, and also brought me a sheep, more in the expectation, I
think, of receiving some Goora nuts in return, than from any regard
for me. At midnight more rain, thunder, and lightning.
June 24.—Cool and cloudy. At ten in the morning halted at the
village of Boorum, to fill our water-skins, and afterwards travelled
through a thick wood, where we saw a number of karigums and
elephants: the karigum is a species of antelope, of the largest size,
as high as a full grown mule. At sunset we pitched our tents in the
woods. The night was extremely boisterous, with rain, thunder and
lightning, and violent squalls of wind; and my tent being blown down,
the baggage was drenched with water.
June 25.—Next morning we continued our route through a thick
wood, and halted at Joba during the heat of the day, when I had my
baggage dried in the sun. We still travelled through a thick wood,
and at seven in the evening encamped at a village called Gorbua.
Rain, thunder, and lightning, all night.
June 26.—Cloudy, with rain. At ten in the morning I left Gorbua, or
“the strong town,” as it is ironically called in the Bornouese language,
from being enclosed with matting. Our road, still winding and woody,
led through the Bede territory; and at sunset we reached Guba, a
small town on the south bank of the Yeou, within the dominions of
Bornou.
June 27.—The forenoon was rainy, which obliged us to remain at
Guba till one in the afternoon; when the weather clearing up, we
loaded the camels, and crossing to the north bank of the channel of
the river, which was now dry, we travelled east by south to the town
of Muznee, where we halted for the night.
June 28.—Cloudy, with rain. We travelled eastward along a
crooked path, full of holes, and overgrown with brushwood, and took
up our abode for the night at the town of Redwa. An officer of the
sultan of Bornou was here, collecting his master’s dues, and sent me
milk, onions, and six fowls; and I presented him, in return, with fifteen
Goora nuts.
June 29.—After travelling east by north, we halted at noon at
Kukabonee, or “wood and fish,” a large town on the south bank of
the Yeou. We next passed Magawin, and a number of other villages
and towns on the banks of the river, which we had not visited before,
when we accompanied the sheikh last year.
June 30.—Cool and cloudy. We halted at ten in the morning at
Dungamee, in consequence of heavy rain with thunder and lightning,
which continued without intermission all day.
July 1.—Clear. The weather was hot and sultry. At sunset we
arrived at Mugabee. I shot at a hippopotamus which was swimming
in a lake, of which there are many in this part of the country; I
seemed to hit it, but it quickly disappeared.
July 2.—Stopped for the day to allow the camels to have food and
rest.
July 3.—Between Gateramaran and Mugabee we met Malam
Fanamee, the governor of Munga, who had been to Kouka on a visit
to the sheikh. He was a dirty looking old man, preceded by a
drummer beating a drum, and attended by a parcel of ragged
followers, armed with bows and spears. We encamped at night in a
wood.
July 4.—At mid-day we halted on the banks of the Yeou: in the
afternoon there was thunder, lightning, and rain. A dealer in fish, who
had joined our party, solicited me in vain to pursue a route through a
town named Sucko, where he was going, promising me a sheep,
with plenty of milk, as an inducement. We passed another night in
the woods.
July 5.—Clear and cool. At ten in the morning we halted and filled
our water-skins, and I here shot a hare and two Guinea-fowls. About
an hour after starting we had heavy squalls of wind, with thunder and
rain: the storm was so violent that the camels lay down with their
burdens, and my horse would neither move forward, nor face the
storm in spite of all I could do. It was an hour before we were able to
resume our journey, and at eight in the evening we encamped in the
woods. The dangers of the road being past, my two fellow travellers,
the merchants before mentioned, left me at midnight on account of
the want of water.
July 6.—To-day I shot a fine male mohur, or beautiful red and
white antelope; a female only of which species I had once shot at
Woodie. At noon we took shelter under the walls of Borgee from
heavy squalls of wind and sand, but without rain. At sunset we
encamped near a well where there had been a great fall of rain, and
all the hollows were filled with water. To roast our mohur a large fire
was kindled in a hole made in the sand, on which it was placed, and
then covered over with hot embers; but, in the morning, to our great
disappointment, nothing remained of our prize but the naked
skeleton.
July 7.—At noon we halted at the wells of Barta, and encamped at
night at the wells and town of Calawawa.
July 8.—At eight in the morning I returned to Kouka: Major
Denham was absent on a journey round the east side of the Tchad.
Hillman, the naval carpenter, was busily employed in finishing a
covered cart, to be used as a carriage or conveyance for the
sheikh’s wives: the workmanship, considering his materials, reflected
the greatest credit on his ingenuity; the wheels were hooped with
iron, and it was extremely strong, though neither light nor handsome.
July 9.—In the afternoon I waited on the sheikh, who was very
kind in his inquiries after my health, and expressed much regret at
Dr. Oudney’s death.
July 10.—To-day the sheikh sent me three pairs of slippers, two
loaves of sugar, and a supply of coffee; and two days afterwards a
sheep, two bags of wheat, and a jar of honey.
APPENDIX.
TRANSLATIONS FROM THE ARABIC, OF VARIOUS LETTERS AND
DOCUMENTS, BROUGHT FROM BORNOU AND SOUDAN BY
MAJOR DENHAM AND CAPTAIN CLAPPERTON.
BY A. SALAME, ESQUIRE.
No. I.
Translation of a Letter from the Sheikh Mohammed El Kanemy,

Chieftain of Bornou, in the Interior of Africa, to his Most Excellent
Majesty King George the Fourth. Brought by Major Denham.
“Praise be to God, and blessings and peace be unto the Apostle

of God (Mohammed). From the servant of the High God, Mohammed
El Ameen ben Mohammed El Kanemy,
“To the pre-eminent above his equals, and the respected among
his inferiors, the great King of the English, salutation be to him from
us:
“Whereas your messengers, the travellers through the earth, for
the purpose, as they state, of seeing and knowing its marvellous
things, have come to us, we welcomed them, and paid attention to
their arrival, in consequence of what we heard of your intercourse
with the Mùslemeen, and the establishment of your friendly relations
between you and their kings, since the time of your and their fathers
and grandfathers (ancestors).
“We have thus regarded that friendship, and behaved to them
according to its merits, as much as God the Omnipotent enabled us.
They communicated your compliments to us, and that which you
stated in your letter, that you would not object, if we should be in
want of any thing from your country, was made known to us; and we
felt thankful to you for this (offer) on your part.
“They are now returning to you, after having accomplished their
wishes; but one of them, whose period of life was ended, died. This
was the physician; and an excellent and wise man he was.
“The Rayes Khaleel (travelling name of Major Denham) desired of
us permission, that merchants seeking for elephant-teeth, ostrich
feathers, and other such things, that are not to be found in the
country of the English, might come among us. We told him that our
country, as he himself has known and seen its state, does not suit
any heavy (rich) traveller, who may possess great wealth. But if a
few light persons (small capitalists), as four or five only, with little
merchandize, would come, there will be no harm. This is the utmost
that we can give him permission for; and more than this number
must not come. If you should wish to send any one from your part to
this country again, it would be best to send Rayes Khaleel; for he
knows the people and the country, and became as one of the
inhabitants.
“The few things that we are in want of are noted down in a
separate paper, which we forward to you.
“Write to the consul at Tripoli, and to that at Cairo, desiring them,
if any of our servants or people should go to them for any affair,
either on land or at sea, to assist them, and do for them according to
their desire. And peace be with you.
“Dated on the evening of Saturday, the middle of the month
Fledja, 1239 of Hejra (corresponding to August 1824).
“Sealed. The will of God be done, and in God hath his faith, his
slave Mohammed El Ameen ben Mohammed El Kanemy.”
No. II.
Translation of a Letter from an African Chieftain (Bello) of Soudan, to

his Majesty King George the Fourth. Brought by Mr. Clapperton.
“In the name of God, the merciful and the clement. May God bless
our favourite Prophet Mohammed, and those who follow his sound
doctrine.
“To the head of the Christian nation, the honoured and the
beloved among the English people, George the Fourth, King of Great
Britain;
“Praise be to God, who inspires, and peace be unto those who
follow, the right path:
“Your Majesty’s servant, Ra-yes-Abd-Allah, (Mr. Clapperton’s
travelling name,) came to us, and we found him a very intelligent and
wise man; representing in every respect your greatness, wisdom,
dignity, clemency, and penetration.
“When the time of his departure came, he requested us to form a
friendly relation, and correspond with you, and to prohibit the
exportation of slaves by our merchants to Ata-gher, Dahomi, and
Ashantee. We agreed with him upon this, on account of the good
which will result from it, both to you and to us; and that a vessel of
yours is to come to the harbour of Racka with two cannons, and the
quantities of powder, shot, &c. which they require; as also, a number
of muskets. We will then send our officer to arrange and settle every
thing with your consul, and fix a certain period for the arrival of your
merchant ships; and when they come, they may traffic and deal with
our merchants.
“Then after their return, the consul may reside in that harbour (viz.
Racka), as protector, in company with our agent there, if God be
pleased.”
“Dated 1st of Rhamadan, 1239 of Hejra.” 18th April, 1824.
No. III.
A Letter from Yousuf, Pasha of Tripoli, to the Sheikh of Bornou.
“Praise be to God, and prayers be unto him who was the last of
the Prophets (Mohammed).
“To the learned and accomplished, the virtuous Iman, the jealous
and zealous defender of the Mohammedan faith, our true friend the
Sheikh Mohammed El Kanemy, Lord of the country of Barnooh[66],
and its dependencies, whom may God protect and dignify, and
prolong his life long in happiness and felicity. Peace be unto you,
and the mercy and blessings of God be upon you, as long as the
inhabitants of the world shall exist.
“It follows, my Lord, subsequent to the due inquiry we make after
your health, which may God preserve, that your esteemed letter has
reached us, and we became acquainted with its contents. You
informed us that our beloved son, Aba Bak’r Ben Khalloom, arrived
in your presence, in company with some persons of the English
nation, our friends; and that you received them with extreme
kindness, and showed them all the marvellous things that your
country contains, and made them see all the extraordinary rivers and
lakes that surround it; and that you behaved to them as becoming
your high station, and indicating your esteem and regard towards us.
May God reward you for all this kindness, and protect you from all
evils. This kind treatment was our sanguine expectation, and indeed
we were already sure of it, from what we knew of the true friendship
and amity established between us.
“What we have now to acquaint you with, is to request that you
will continue your protection and assistance to the said English
travellers (though we doubt not you do not need this additional
recommendation), and cause them to proceed to the country of
Soudan, to behold its marvellous things, and traverse the seas
(lakes or rivers), and deserts therein. This being the proper desire of
the great King of the English himself, we beg of you to use your
utmost endeavours, as far as lies in your power, in their safe arrival
at the country of Soudan, accompanied either by letters of
recommendation, or by troops and guards, in order that they may
obtain the accomplishment of their wishes, and return to us safe and
unhurt; and whatever kindness you may do to them, it is done to us.
Resolve therefore, and exert yourself, as we are confident of your
goodness, and let them see all the places which they wish to visit.
“At the end there will be a splendid present, befitting your high
rank, sent to you through us, consisting of various rare and elegant
articles of value; for the delivery of which, unto your hands, we
pledge ourselves.
“This is all that we have to say at present, and if any affair should
occur to you in this country, let us know. And peace be unto you.
“Your friend,
(Signed) “YOUSUF PASHA .”
(Dated) “28th of Sha-wal, 1238 of Hejra;”
corresponding to August, 1823.
FOOTNOTES:
[66]Note. This is the proper name of Bornou. A. S.

No. IV.
A Letter from the before named Pasha of Tripoli to Aba Bak’r ben
Khalloom, at Bornou.
“We received your letter, and comprehended all that you stated to
us. We were glad to hear that you, and our friends, the English
travellers, with whom we sent you as guide and conductor, had
arrived at Barnooh in safety; and that you were kindly received by
our friend, my Lord, the Sheikh Mohammed El Kanemy, who
immediately allowed the travellers to inspect all the deserts, and
seas, lakes and rivers, that are in his country. May God reward him
for this act of kindness. We have written to thank him for his laudable
behaviour; and we pray to God to enable us to show him equal
kindness in return.
“With regard to the persons of the different tribes, who were
obstinate and disobedient to you on the road, they have been
apprehended, and taken and punished one by one.
“As long as the English travellers remain at Barnooh, you have to
attend, and be with them wherever they go, till they shall have
obtained their wishes, and accomplished their object; and when they
desire to return, you may accompany and come with them as you
went. If this letter should reach you before you leave Barnooh, you
must stay with them, as above stated; if it reach you while you are on
the road homewards, you must return to Barnooh immediately, and
only send us the slave you have with you; and if you should arrive at
Fezzan before this letter reaches you, you may then send your
brother to Barnooh, to stay with them instead of you; for we only sent
you on their account, for the purpose of facilitating their proceeding,
and all their affairs. It is, therefore, impossible that you should leave
or part with them, but in this manner; and we are sure that, to a
person like you, there is no need to add any stronger words,
especially as you know that they are in our honour, and under our
protection, both in their going and returning in safety; which is the
accomplishment of our wishes. And may you live in happiness and
peace.
(Signed) “YOUSUF PASHA .”
(Dated) “2d of Ze-el-ka’da, 1238;”
corresponding to August, 1823.
No. V.
A Letter from the Sheikh of Bornou to the Sultan of Kanou.
“Praise be to God, and prayers and peace be unto the Apostle of

God (Mohammed).
“From the slave of the high God, Mohammed El-ameen ben
Mohammed El-kanemy, to the head of his land and the leader of his
people, the learned Mohammed Daboo, lord and master of Kanou:
Perfect peace, and the mercy and blessings of God, be unto you.
“Hence, the bearer, who is going to you, is our friend Mohammed
El-wardy, in whose company he has some Englishmen; who came to
the land of Soodan for the purpose of seeing and delighting
themselves with the wonders it contains, and to examine and see the
lakes and rivers, and forests, and deserts therein. They have been
sent by their king for this purpose.
“Between their nation and the Mooslemeen, there have existed,
since the times of their fathers and great grandfathers (ancestors),
treaties of religious amity and friendship, special to themselves out of
all the other nations that have erred, and are at variance with the
doctrine of Aboo Hanifa[67]. There never was between them and the
Mooslemeen any dispute; and whenever war is declared by the other
Christians against the Mooslemeen, they are always ready to help
us, as it has happened in the great assistance they gave to our
nation when they delivered Egypt from the hands of the French.
They have, therefore, continually penetrated into the countries of the
Mooslemeen, and travelled where-ever they pleased with confidence
and trust, and without being either molested or hurt. They are, as it is
stated, descendants of the ancient Greek emperor Heraclius, who
received and esteemed the letter sent to him from the Apostle of
God (Mohammed), whom may God bless, by Dahi-yah El-kalbee,
containing his exhortation to him to embrace the Moosleman faith;
and who, on receiving that sacred epistle, preserved it in a gold
case,—though it is stated, in the books of history, that he did not
become a Mooslem.
“Thus, if God permit them to reach you in safety, be attentive to
them, and send guards to conduct them to the country of Kashna,
safe and unhurt; for they are at the mercy of God, and at the honour
of his Apostle; and you are well aware of the Alcoraanic sayings
upon the subject of the observance of honour. And peace be with
you.”
Dated “Wednesday, the 6th day of Rabee-ul-thani, 1239,”
(Corresponding to January, 1824.)
FOOTNOTES:
[67]Aboo Hanifa, or Imam Kanafee, was one of the four great

imams or high priests, founders of the four orthodox rites of
Mohammedanism; and whose doctrine, it seems, is followed by
these people. A. S.
No. VI.
A Letter from the Sheikh of Bornou to Mohammed Bello, Sultan of

Hoossa.
“Praise be to God, and prayers and peace be unto the Apostle of

God, (Mohammed).
“To the honoured and accomplished, the virtuous and munificent,
the pattern of goodness and the standard of benevolence, head of
the Soodanic kingdom, and ruler of the country of Hoossa, our
friend, the learned Mohammed Bello, son of the intelligent sheikh
Ossman, whose soul may God shelter with the clouds of mercy and
peace.
“Our kind salutation, accompanied with affection as strong as the
odour of musk, and as perpetual as the movement of the globe, and
with the mercy and blessings of God, be unto you.
“Hence, the cause of writing this letter and the purpose of its lines,
is to acquaint you that the bearers are English travellers; whose
nation, out of all the other Christians, has maintained with the
Mooslemeen uninterrupted treaties of religious amity and friendship,
established since ancient periods, which they inherited from their
forefathers and ancestors; and, on this account, they penetrate into
the Mooslemeen countries whenever they please, and traverse all
provinces and lands, in confidence and trust, without fear. They
came to our country, sent to us by our virtuous and accomplished
friend, the Lord Yousuf Pashá, master of Tripoli, to see and delight
themselves with the wonders of the land of Soodan, and to become
acquainted with its rarities, as lakes, rivers, and forests (or gardens);
equal to which are seldom seen in any other countries.
“After having accomplished their wishes, in seeing all the things
that the land of Barnooh and its environs contained, they felt anxious
to visit your country from what they heard of the innumerable
wonders therein. I have, therefore, permitted them to proceed on
their journey, accompanying them with letters which explain their
object.
“You are well aware of what is stated in the Alcoraanic sayings
upon the subject of the observance of honour, dictated by our Lord,
the Apostle of God; and that the true Mooslemeen have always
avoided shedding the blood of Christians, and assisted and
protected them with their own honour. Be then attentive to these
travellers, and cast them not into the corners of neglect; let no one
hurt them, either by words or deeds, nor interrupt them with any
injurious behaviour: but let them return to us, safe, content, and
satisfied, as they went from us to you; and may the high God bestow
upon you the best reward for your treatment to them, and insure to
us and to you the path of the righteous for our conduct in this life.
“Our salutation may be given to all who are about you, and to
those who are related to you in general. And peace be unto you.

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Vanders Human Physiology The

Mechanisms of Body Function 13th

SECTION A: SKELETAL MUSCLE

9.2 Molecular Mechanisms of Skeletal Muscle Contraction

9.3 Mechanics of Single­Fiber Contraction

9.4 Skeletal Muscle Energy Metabolism

9.5 Types of Skeletal Muscle Fibers

9.6 Whole­Muscle Contraction

9.7 Skeletal Muscle Disorders

Suggestion for class discussion: World­class athletes and muscle hypertrophy/changes in

I. Three types of muscle fibers (Fig. 9‐1)

SECTION A REVIEW QUESTIONS

1. List the three types of muscle cells and their locations.

4. Describe the four steps of one cross‐bridge cycle.

10. Describe the structure and function of the transverse tubules.

12. Define a motor unit and describe its structure.

14. What is an end‐plate potential, and what ions produce it?

16. Describe isometric, concentric, and eccentric contractions.

21. What is the function of creatine phosphate in skeletal muscle contraction?

23. List the factors responsible for skeletal muscle fatigue.

SECTION B: SMOOTH AND CARDIAC MUSCLE

9.8 Structure of Smooth Muscle

9.9 Smooth Muscle Contraction and Its Control

9.10 Cardiac Muscle

SECTION B REVIEW QUESTIONS

5. What effect does a pacemaker potential have on a smooth muscle cell?

8. Describe the differences between single‐unit and multiunit smooth muscle.

10. Explain why cardiac muscle cannot undergo tetanic contractions.

ANSWERS TO TEST QUESTIONS

9­1 a A single skeletal muscle fiber, or cell, is composed of many myofibrils.

ANSWERS TO QUANTITATIVE AND THOUGHT QUESTIONS

9­5 Moderate tension—for example, 50 percent of maximal tension—is accomplished by recruiting

ANSWERS TO GENERAL PRINCIPLES ASSESSMENTS

Translation of a Letter from the Sheikh Mohammed El Kanemy,

“Praise be to God, and blessings and peace be unto the Apostle

Translation of a Letter from an African Chieftain (Bello) of Soudan, to

A Letter from Yousuf, Pasha of Tripoli, to the Sheikh of Bornou.

[66]Note. This is the proper name of Bornou. A. S.

A Letter from the Sheikh of Bornou to the Sultan of Kanou.

“Praise be to God, and prayers and peace be unto the Apostle of

[67]Aboo Hanifa, or Imam Kanafee, was one of the four great

A Letter from the Sheikh of Bornou to Mohammed Bello, Sultan of

“Praise be to God, and prayers and peace be unto the Apostle of

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9.3 Mechanics of SingleFiber Contraction

9.6 WholeMuscle Contraction

Suggestion for class discussion: Worldclass athletes and muscle hypertrophy/changes in

91 a A single skeletal muscle fiber, or cell, is composed of many myofibrils.

95 Moderate tension—for example, 50 percent of maximal tension—is accomplished by recruiting