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PHYSIOLOGY
THEME II
CHAPTER OUTLINE
To Describe
To Define
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THE JOINT SYSTEM
The need for strength requires bones to be
rigid. However, movement would be almost impos-
sible if our skeletons were solid bone. Nature has
solved this problem in vertebrates, including humans,
by casting the skeleton into many bones and creating
joints where they intersect. Joints are custom-formed
to serve the functional needs of the limbs that contain
them.
They are held together by fibrous tissue (cap-
sule and ligaments) and continuously lubricated to
offset friction. In this way joints permit motions, which
grant us humans a very complex repertoire of move-
ment.
The word articulation and joint are used syn- Figure - Ligaments of the knee joint.
onymously to refer to those structural arrangements
that connect two or more bones. Although most joints
permit at least some movement between the bones DEVELOPMENT
they connect, this is not essential for a connecting
structure to be called a joint. The function of some Bone and joint both develop from the middle
joints is to allow the joined structures to grow until layer of tissue in the embryo. Circumscribed conden-
they in turn become as solid as the bones they con- sation of cells in this mesoderm become chondrified
nect. (turned into cartilage) and finally ossified (turned into
Where joints are immovable, as in the articu- bone) to form the bones of the skeleton. The inter-
lations between the bones of the growing skull, the vening non-condensed portions of tissue develop
adjacent margins of the bone are separated merely into joints.
by a thin layer of fibrous tissue. Where slight move- As soon as the joint cavity appears during de-
ment but great strength is required, the joint surfaces velopment, it contains watery fluid. The tissue sur-
are united by tough elastic fibrocartilages, as in the rounding the original mesodermal cellular core forms
joints between the vertebral bodies. In freely movable fibrous sheaths for the developing bones, which
joints, such as the shoulder or knee, the surfaces are continues between their ends as the capsules of the
completely separated and the bones expanded for joints. Ligaments develop both in these capsules and
greater convenience of mutual connection. They are as derivations from tendons surrounding the joint.
covered by cartilage and enveloped by fibrous tissue After the joint cavity is established during the third
capsules. month of gestation, the muscles that move the joint
begin to contract. This movement enhances nutrition
of the articular car
ARTICULAR CARTILAGE tilage and prevents fusions of the apposed
joint surfaces. Early restriction of joint motion can
Articular cartilage is of a type called hyaline result in permanent loss of the joint cavity, whereas
cartilage. In contrast to bone, it is easily cut. It is de- later restriction can lead to abnormalities of the soft
formable under pressure, but elastic so that it quickly tissues associated with the joint.
recovers its shape. These properties are important Because normal positioning of the fetus in the
for its function under conditions of loading (bearing uterus permits a fair degree of movement of the up-
weight). A membrane lines the interior of such joints. per limbs but restricts the legswhich are folded and
This synovial membrane secretes a thick vis- pressed firmly against the bodythe lower limbs are
cous liquid that lubricates the joint. It also regulates more vulnerable to congenital joint deformity. In sev-
protein and electrolyte metabolism in the joint and re- eral of the movable joints, a portion of the mesoder-
moves waste from the joint. Joints are strengthened mal tissue that originally existed between the ends
by ligaments, strong fibrous bands that connect the of the bones persists and forms an articular disc. An
bones that form the joint.
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example of this is the menisci of the knee joint. SYNOVIAL JOINTS
MECHANICS
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THE BONY SYSTEM Osteoblasts make bone, osteoclasts remodel bone,
osteocytes govern its metabolism, and a group of
The unique tissue that is bone is found only in uncharacterized basic structures termed undifferenti-
vertebratesanimals with backbones. ated mesenchymal cells stand ready to evolve into
Newborn infants have approximately 350 individual -blasts, -clasts, or -cytes if and when needed. With
bones. By fusion and rearrangement during growth, the exception of dental enamel, bone is the hardest
they develop into their adult counterparts. The adult tissue in the body. Now lets see how bone renews
skeleton (Greek: skeletos, meaning dried up) usually itself.
contains 206 bones, although some people are born
with either an extra pair of ribs or one pair less.
Bone consists of living cells held in a hard As in any other mammal, human bones are
intercellular material. This material is composed of preformed in cartilage. This soft tissue model
collagen, a fibrous protein similar in structure and or- is gradually converted into bone tissue by a bony
ganization to ligaments, tendons, and skin. This col- change called osteogenesis. A skeletal outline can be
lagenous matrix is mineralized, mainly with calcium seen in the human embryo at approximately the fifth
and phosphorus. week of pregnancy. Ossification of bone continues
The intimate blending of the hard and resilient until almost 25 years of age, at which time the carti-
components in bone makes it almost equally resist- lage is completely replaced by bone, ending further
ant to compression and tension. This is in contrast transformation. A few of the bones of our bodies are
to most of the structural materials that we commonly not preformed in cartilage. These membrane bones
use, which are usually better in one respect than an- form in condensed fibrous tissue. They include the
other. The two-phased materials used in bone give it bones of the skull and the clavicle (collarbone).
hardness, rigidity, and flexibility. This is similar to oth- Long bones grow at each end through a growth
er two-phased materials, such as fibreglass or bam- plate, or physis, as well as from a thin membrane
boo. Because an element of viscous flow is present, called the periosteum, which surrounds the shaft of
bone is stiffer during rapid deformation than during the bone. Cells from this layer begin to form bone
slow deformation. around the middle of a cartilage shaft and continue
The remaining portion of bone is composed this process during growth, which allows the bone to
of collagen, the elastic protein that makes glue when increase in diameter as well as in length. At the ends
bones are boiled. Fibers of collagen studded with of bones, cartilage cells are destroyed and replaced
crystals wind themselves around each other like by bone tissue. These newly deposited cells push the
strands of rope fibers. Remove the mineral in bone growth plate farther from the center of the shaft until
and the bone becomes so pliable that it can be tied in the body has attained its full height, which occurs at
a knot. approximately 18 years in men and two years earlier
The skeleton demonstrates the use of state- in women. During our lifetime, we actually have three
of-the-art engineering principles to provide the great- skeletonsthe first one is made of cartilage, the sec-
est strength with the least material. It closely resem- ond of an immature fibrous type bone, and the third
bles coral in structure, and like such meshworks it is of adult bone.
both sturdy and delicate. Numerous metabolic influences affect bone
The femur, or thigh bone, is the longest and growth. Essential monitoring and manufacturing roles
strongest of all human bones. It may be called on in the development of the bony skeleton are played
to resist a compressive force of more than 1,000 by the availability of minerals such as phosphorus
pounds per cubic inch during walking. What is even and calcium, vitamins A, D, and C, as well as by the
more remarkable is that, unlike nonliving materials secretions of many endocrine glands, including the
such as concrete or steel, bone remodels itself under thyroid, parathyroid, pituitary, adrenals, and gonads.
stress. It does this through a complex system of cel- The tissues of the body constantly renew
lular communication that is electrical in nature. themselves. This process is so rapid and so continu-
Like any well-recognized factory, bone di- ous that you hardly have any single cell in your body
vides labour between its four major types of cells. that you had three years ago. Renewal in soft tissues
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occurs largely at a molecular level. Bony change oc- these ions by the intestine. Vitamin D is required in
curs by replacing tissue; it is somewhat like urban this metabolic scheme. Calcitonin is a body chemi-
renewal in that removal of the old bone by resorption cal that inhibits bone resorption. Renovation of bone
must precede new bone deposition. Remodelling is depends on two of these control loops with negative
greatest during growth and declines until approxi- feedback. Remodelling under Wolffs law may be re-
mately age 35. After the age of 35, remodelling re- lated to the piezoelectric properties of bone crystals,
mains essentially unchanged. After the fourth decade which involve the generation of electricity by crystal-
of life, resorption of bone is greater than formation, line structures under stress. Tension on a bone sets
and a loss of 5 percent to 10 percent of bone mass up a negative electrical pole (anode), whereas com-
occurs with each passing decade. This works out to pression sets up a positive pole (cathode). This proc-
a loss of approximately 4 grams of calcium from the ess also plays an important role in bone healing.
skeleton each year. A number of hormones also play significant
The processes of remodelling, growth, and roles in the regulation of bone growth and metab-
development are under the control of the cells of olism. These include estrogens, testosterone, thy-
bone. Osteoclasts secrete enzymes that dissolve roid hormone, adrenal gland hormones, insulin, and
bone, and osteoblasts elaborate a protein-like sub- growth hormone.
stance, called osteoid, which is then mineralized. It
is thought that osteocytes, the mainstay cell of ma-
ture bone, are also metabolically active and can re-
sorb and even re-form bone on demand. To keep the THE MUSCLE PHYSIOLOGY
delicate mineral chemistry of the body in balance,
bone acts as the bodys major storehouse for miner- The number of muscle fibers in the body is
als. Approximately half of bones volume and up to 75 determined at birth. In this respect, muscle is similar
percent of its weight consists of deposited minerals. to nervous tissue. The number of fibers does not mul-
Almost 90 percent of the calcium and phosphorus in tiply, but individual fibers do enlarge with work. This
the body is found in the bones. If spread out, the crys- accounts for the size and strength of the muscles of a
tal surface exposed to the body fluids in the walls of weight lifter contrasted with those of an inactive desk
the skeletal system of an average man would cover clerk, even though their muscles contain the same
1,500 to 5,000 square meters. This is approximately number of fibers. Theoretically, the maximum energy
the size of a football field and provides an enormous output of a man is approximately 6 hp. The highest
surface area across which chemical exchanges of recorded output to date is 4.5 hp; 0.5 hp can be sus-
minerals can occur. tained almost indefinitely.
Julius Wolff, who was a professor of ortho- Muscle fibers can be short or long, but their
paedics in Berlin, formulated his famous law of bone diameter is always less than their length (most fib-
modelling late in the nineteenth century. Wolffs law ers measure from 0.01 mm to 0.1 mm across). Many
holds that the physical stresses placed on bone ini- individual fibers make up each muscle. There are ap-
tiate accommodative changes in its structure and proximately 650 skeletal muscles in the body (about
form. It would seem that small electrical currents are three times as many muscles as bones), containing
generated from pressure in bone, and these charges about 250,000,000 striated muscle fibers. In the eye
stimulate resorption or production in response to the these fibers have a diameter of 20 microns (one mi-
stress imposed. cron is one millionth of a meter). The fibers of the
The human skeleton not only serves to store limbs range in size from 10 to 100 microns, being
minerals but also acts as the reservoir that controls largest in male athletes.
the equilibrium of calcium and phosphate in the Each muscle obeys the same all-or-none
blood stream. These elements are vital for cardiac principle as the nerve fiber that controls iteither it
function, muscle contractility, cell membrane integrity contracts or it doesnt. Muscle works by converting
and permeability, neuromuscular function, and blood chemical energy to mechanical energy. Less than
clotting. Bone governs the body concentration of cal- half of the potential energy available is converted,
cium and phosphorus through hormonal regulation and the remainder is lost as heat. This is why the
by parathyroid hormone and calcitonin. body temperature rises when you perform strenuous
This is controlled through feedback loops work. Even at rest, the heat produced during the rela-
that hormonally link the excretion of calcium and tively efficient conversion of chemical to mechanical
phosphorus by the kidney with the absorption of energy in muscle maintains body temperature in
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animals exposed to the cold.
The muscular movements associated with
shivering liberate heat so it is not wasted, as it is in
most man-made machines. Sometimes enough heat
is produced to boil a quart of water. Shivering may be
better than a heating pad or exercise when it comes
to warming
the body. In some cases of modest hypothermia, heat
from an external source may actually be counterpro-
ductive because it inhibits the shivering response.
STRUCTURE
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(muscle cellular units) binding them together and to The Growth and Development of Muscle
perimysium. Within this fibrous envelope the muscle
cells are bathed in a fluid that contains the chemicals Muscle develops from a group of cells called
needed for contraction and relaxation. Access from mesenchyme (middle layer). The outer membranes
the general circulatory system is through microscop- of these cells fuse at contact points during embry-
ic tubules. Blood vessels and nerves also enter this ologic development, forming a single unit enclosed
system. in a continuous membrane called a myoblast (rnyo
Tiny spindles of highly specialized muscle = muscle, blast = formative cell). This early organi-
cells served by nerves that regulate position are found zation results in a so-called myotube, which in turn
throughout postural muscles. These muscle spindles develops into an immature muscle cell that may con-
register muscular tension so that fine postural adjust- tain hundreds of nuclei, one from each of the original
ments can be made by the body where and when cells. Further growth produces a mature muscle cell,
necessary. Another type of tension regulator is the or fiber, a long cylinder that measures 1/500 of an
Golgi tendon organ receptor found in tendons, which inch in diameter and several inches in length. Disrup-
supplies the nervous system with information about tion of development at any point along this path can
muscle tension to prevent extremely forceful contrac- lead to a congenital disease of muscle.
tions that might separate tendon from bone.
In spite of variations in size and shape, all
muscles have essentially the same structure. Myofi-
brils consist of muscle filaments arranged in a repeat-
ing pattern. Each unit of this pattern is called a sar-
comere. Each sarcomere contains several types of
filamentsthick ones containing the protein myosin,
and thin ones containing the protein actin.
About 100,000 myosin molecules laid side by
side would form a ribbon 1 mm wide. Actin molecules
are half as thick. The serial alignment of sarcomeres
in adjacent myofibrils gives muscle fibers their stri-
ated (striped) appearance.
NERVE CONTROL
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REPRODUCTION filaments alternate. Various lines, bands, and zones
have been noted in the pattern and labelled (Z, I, A,
Muscle cells pay for their specialized con- M, H) for identification during microscopic examina-
tractile ability. One of the things they cannot do is tion.
divide in order to reproduce themselves in the con- As the muscle fibril contracts, the thin fila-
ventional manner of most cells. In humans and most ments slide between the thick filaments. Molecular
other mammals, the number of muscle cells you have arms on the thick (myosin) filament engage specific
at birth is the number you have to work with as you sites on the thin (actin) filament and by a sort of ratch-
grow. et action shorten the fiber in contraction.
Muscle mass is increased through growth
and exercise, by increasing the size of the existing
cells. However, specialized satellite cells can regen-
erate muscle cells that have been damaged through
an intricate process of assembling structures already
floating in the cell fluid. The muscle cell undergoes
necrosis (death) and the satellite cell directs regen-
eration, which includes splitting of the nucleus and
reduplication of the membrane containing the cell
(the basal lamina). Eventually, however, the normal
attrition that results from aging catches up with this
process. The weight of some muscles may decrease
by as much as 30 percent between the ages of 35
and 75.
A number of things can accelerate loss of
muscle. Stimulation to its muscle is cut off when a
nerve is destroyed, as with polio, Lou Gehrigs dis-
ease (amyotrophic lateral sclerosis, or ALS), or trau-
ma, and the muscle fiber becomes inactive, shrinks,
and eventually is replaced by fat and connective
tissue. The reverse situation results in a highly ac-
tive muscle that increases in size and power. In this
sense, muscular development can be looked upon as
a direct product of muscle use.
ORGANIZATION
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ceptors on the muscle, the fiber is akin to a charged more carbohydrate is used, with less reliance on fat.
battery carrying an electrical charge of 0.09 volt. This In maximal, short-duration exercise, ATP is gener-
is called the resting potential of muscle. ated almost exclusively from carbohydrate.
Acetylcholine is a neurohumor (nervous sys- Carbohydrate
tem chemical secretion) that can temporarily change Your muscles dependence on carbohydrate
the molecular structure of the muscle membrane so during exercise is related to carbohydrate availability
that there is a two-way flow of potassium and sodium and your muscles well-developed system for its me-
ions (an ion is an atom that is electrically charged) tabolism. Carbohydrates are ultimately converted to
between the muscle cell and the fluid surrounding it. glucose, a monosaccharide (one-unit sugar) that is
As these ions are exchanged through the transported via your blood to all body tissues. Under
membrane, the electrical balance of the muscle resting conditions, ingested carbohydrate is taken up
fiber changes (depolarizes) as it becomes temporar- by your muscles and liver, and then converted into a
ily discharged. This generates an action potential, an more complex sugar molecule: glycogen.
electrical discharge that spreads over the surface of Glycogen is stored in the cytoplasm until your
the cell, transmitting a message through a tubular cells use it to form ATP. The glycogen stored in your
pathway called the T-tubules to a holding area (the liver is converted back to glucose as needed, then
terminal cistern) within the membrane (sarcoplas- transported by your blood to active tissues, where it
matic reticulum, or SR) that surrounds the innermost is metabolized.
contracting elements, the myofibrils. Liver and muscle glycogen reserves are lim-
Finally, an interchange of calcium ions deep ited and can be depleted unless the diet contains a
within the fibrils initiates contraction. The total time reasonable amount of carbohydrate. Thus, we rely
involved in a single muscle cell twitch, including stim- heavily on dietary sources of starches and sugars
ulation, contraction, and relaxation, is approximately to replenish our carbohydrate reserves, without ad-
0.1 second. equate carbohydrate intake, the muscles and liver
Millions of muscle fibers are activated with can be deprived of their primary energy source.
every body movement. Electrical interactions with
subsequent chemical changes constantly occur as Fat
electricity flows back and forth through the cell mem-
branes and along pathways within the fibers. The Fat provides a sizable amount of energy dur-
pacemakers that orchestrate this activity are the ing prolonged, less intense exercise. Body stores of
nerve signals that release acetylcholine at multitudi- potential energy in the form of fat are substantially
nal neuromuscular junctions. larger than the reserves of carbohydrate. But fat is
less accessible for cellular metabolism because it
must first be reduced from its complex form, triglyc-
eride, to its basic components, glycerol and free fatty
THE ENERGY METABOLISM acids (FFA). Only free fatty acids are used to form
ATP.
Energy Sources Carbohydrate stores in the liver and skeletal
muscle are limited to less than 2,000 kcal of energy,
Foods are composed primarily of carbon, or the equivalent of the energy needed for about 32
hydrogen, oxygen, and in the case of protein ni- km (20 miles) of running. Fat stores, how-ever, gen-
trogen. Molecular bonds in foods are relatively weak erally exceed 70,000 kcal of energy also because
and provide little energy when broken. Consequently, substantially more energy is derived from a given
food is not used directly for cellular operations. Rath- quantity of fat (9 kcal/g) than from the same amount
er, the energy in food molecules bonds is chemically of carbohydrate (4 kcal/g). Nonetheless, the rate of
released within our cells then stored in the form of energy release from fat is too slow to meet all of the
a high-energy compound called adenosine triphos- energy demands of intense muscular activity.
phate (ATP).
At rest, the energy that your body needs is Protein
derived almost equally from the breakdown of car-
bohydrates (CHO) and fats. Proteins are your bodys Protein can also be used as an energy source,
building blocks, usually providing little energy for cel- but it must first be converted into glucose. In the case
lular function. During mild to severe muscular effort, of severe energy depletion or starvation, protein may
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even be used to generate free fatty acids for cel- phorylation. Through various chemical reactions, a
lular energy. The process by which protein or fat is phosphate group is added to a relatively low-energy
converted into glucose is called gluconeogenesis. compound, adenosine diphosphate (ADP), convert-
The process of converting protein into fatty acids is ing it to adenosine triphosphate (ATP). When these
termed lipogenesis. Protein can supply up to 5% or reactions occur without oxygen, the process is called
10% of the energy needed to sustain prolonged exer- anaerobic metabolism. When these reactions occur
cise. Only the most basic units of protein the ami- with the aid of oxygen, the overall process is called
no acidscan be used for energy. A gram of protein aerobic metabolism, and the aerobic conversion of
yields about 4.1 kcal. ADP to ATP is oxidative phosphorylation
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breakdown (lysis) of glucose. This system is called formula C3H6O3. Lactate is any salt of lactic acid.
the glycolytic system because it involves glycolysis, When lactic acid releases H+, the remaining com-
which is the breakdown of glucose via special glyco- pound joins with Na+ or K+ to form a salt. Anaerobic
lytic enzymes. Glucose accounts for about 99% of all glycolysis produces lactic acid, but it quickly dissoci-
sugars circulating in the blood. Blood glucose comes ates, and the salt lactate is formed. For this reason,
from the digestion of carbohydrate and the break- the terms are often used interchangeably.
down of liver glycogen. Glycogen is synthesized from A muscle fibers rate of energy use during
glucose by a process called glycogenesis. Glycogen exercise can be 200 times greater than at rest. The
is stored in the liver or in muscle until needed. At that ATP-PCr and glycolytic systems alone cannot supply
time, the glycogen is broken down to glucose-1-phos- all the needed energy. Without another energy sys-
phate through the process of glycogenolysis. tem, the exercise capacity might be limited to only a
Before either glucose or glycogen can be few minutes. Then, a third energy system, the oxi-
used to generate energy, it must be converted to a dative system, can provide the necessary energy to
compound called glucose-6-phosphate. Conversion continue any muscular effort from this point on.
of a molecule of glucose requires one molecule of
ATP. In the conversion of glycogen, glucose-6-phos- The Oxidative System: The final system of
phate is formed from glucose- 1-phosphate without cellular energy production is the oxidative system.
this energy expenditure. Glycolysis begins once the This is the most complex of the three energy systems.
glucose-6-phos-phate is formed. The process by which the body disassembles fuels
Glycolysis ultimately produces pyruvic acid. with the aid of oxygen to generate energy is called
This process doesnt require oxygen, but the use cellular respiration. Because oxygen is used, this is
of oxygen determines the fate of the pyruvic acid an aerobic process. This oxidative production of ATP
formed by glycolysis. In the case of the anaerobic occurs within special cell organelles: the mitochon-
glycolysis, without the need for oxygen, the pyruvic dria. In muscles, these are adjacent to the myofibrils
acid is converted to lactic acid. and are also scattered throughout the sarcoplasm.
Glycolysis, which is far more complex than the Muscles need a steady supply of energy to
ATP-PCr system, requires 12 enzymatic re-actions continuously produce the force needed during long-
for the breakdown of glycogen to lactic acid. All these term activity. Unlike anaerobic ATP production, the
enzymes operate within the cells cytoplasm. The net oxidative system has a tremendous energy-yielding
gain from this process is 3 moles (mol) of ATP formed capacity, so aerobic metabolism is the primary meth-
for each mole of glycogen broken down. If glucose od of energy production during endurance events.
is used instead of glycogen, the gain is only 2 mol This places considerable demands on the bodys
of ATP because 1 mol is used for the conversion of ability to deliver oxygen to the active muscles.
glucose to glucose-6-phosphate.
This energy system does not produce large Oxidation of Carbohydrates
amounts of ATP. Despite this limitation, the combined
actions of the ATP-PCr and glycolytic systems allow In carbohydrate metabolism, glycolysis plays
the muscles to generate force even when the oxygen a role in both anaerobic and aerobic ATP production.
supply is limited. These two systems predominate The process of glycolysis is the same whether or not
during the early minutes of high-intensity exercise. oxygen is present. Presence of oxygen determines
Another major limitation of anaerobic glycoly- only the fate of the end product, pyruvic acid. Re-
sis is that it causes an accumulation of lactic acid in call that anaerobic glycolysis produces lactic acid
the muscles and body fluids. In all-out sprint events and only 3 mol of ATP per mole of glycogen. In the
lasting 1 or 2 min, the demands on the glycolytic sys- presence of oxygen, however, the pyruvic acid is
tem are high, and muscle lactic acid levels can in- converted into a compound called acetyl coenzyme
crease from a resting value of about 1 mmol/kg of A (acetyl CoA). Once formed, acetyl CoA enters the
muscle to more than 25 mmol/kg. This acidification Krebs cycle (citric acid cycle), a complex series of
of muscle fibers inhibits further glycogen breakdown chemical reactions that permit the complete oxida-
because it impairs glycolytic enzyme function. In ad- tion of acetyl CoA. At the end of the Krebs cycle, 2
dition, the acid decreases the fibers calcium-binding mol of ATP have been formed, and the substrate (the
capacity and thus may impede muscle contraction. compound upon which the enzymes act, in this case,
Lactic acid and lactate is not the same the original carbohydrate) has been broken down into
compound. Lactic acid is an acid with the chemical carbon dioxide and hydrogen.
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Energy Yield from Carbohydrates: The oxi- ited by the oxygen transport system, so carbohydrate
dative system of energy production can generate up is the preferred fuel during high-intensity exercise.
to 39 molecules of ATP from one molecule of glyco- The advantage of having more carbon in FFA than in
gen. If the process begins with glucose, the net gain glucose is that more acetyl CoA is formed from the
is 38 ATP molecules (recall that one ATP molecule is metabolism of a given amount of fat, so more acetyl
used for conversion to glucose-6-phosphate before CoA enters the Krebs cycle and more electrons are
glycolysis begins). sent to the electron transport chain. This is why fat
Oxidation of Fat metabolism can generate so much more energy than
As noted earlier, fat also contributes lo mus- glucose metabolism.
cles energy needs. Muscle and liver glycogen stores
may be able to provide only 1,200 lo 2,000 kcal of Protein Metabolism
energy, but the fat stored inside muscle fibers and
in fat cells can supply at least 70,000 to 75,000 kcal, As noted earlier, carbohydrates and fatty ac-
even in a lean adult. ids are the bodies preferred fuels. But proteins, or
Although many chemical compounds (such rather the amino acids that form them, are also used.
as triglycerides. phospholipids, and cholesterol) are Some amino acids can be converted into
classified as fats, only triglycerides are major energy glucose (by gluconeogenesis). Alternatively, some
sources. Triglycerides are stored in fat cells and be- can be converted into various intermediates of oxida-
tween and within skeletal muscle fibers to be used tive metabolism (such as pyruvate or acetyl CoA) to
for energy, a triglyceride must be broken down to its enter the oxidative process.
basic units: one molecule of glycerol and three mol- Proteins energy yield is not as easily deter-
ecules of free fatty acids (FFA). This process is called mined as that of carbohydrate or fat because pro-
lipolysis, and it is carried out by enzymes known as tein also contains nitrogen. When amino acids are
lipases. The FFA is the primary energy source. Once catabolized, some of the released nitrogen is used
freed from glycerol, FFA can enter the blood and be to form new amino acids, but the remaining nitro-
transported throughout the body, entering muscle fib- gen cannot be oxidized by the body. Instead it is
ers by diffusion. Their rate of entry into the muscle converted into urea and then excreted, primarily in
fibers depends on the concentration gradient. In- the urine. This conversion requires the use of ATP, so
creasing the FFA concentration in the blood drives some energy is spent in this process.
them into the muscle fibers. To accurately assess the rate of protein
metabolism, the amount of nitrogen being eliminat-
Oxidation ed from the body must be determined. These meas-
urements require urine collection for 12-to 24-h pe-
Although the various FFA in the body differ riods, clearly a time-consuming process. Because
structurally, their metabolism is essentially the same. the healthy body uses little protein during rest and
On entering the muscle fiber, FFA are enzymatically exercise (usually far less than 5% to 10% of total
activated with energy from ATP, preparing them for energy expended), estimates of energy expenditure
catabolism (breakdown) within the mitochondria. generally ignore protein metabolism.
This enzymatic catabolism of fat by the mito-
chondria is termed (beta) oxidation.
From this point on, fat metabolism follows the same
path as carbohydrate metabolism. Acetyl CoA formed THE ENDOCRINE SYSTEM
by oxidation enters the Krebs cycle. As in glucose
metabolism, the by-products of FFA oxidation are ATP The endocrine system works in parallel with
H2O, and CO2. However, the complete combustion the nervous system to control growth and maturation
of an FFA molecule requires more Oxygen because along with homeostasis. It is a collection of glands
an FFA molecule contains considerably more carbon that secrete chemical messages we call hormones.
than a glucose molecule. These signals are passed through the blood
Although fat provides more kilocalories of to arrive at a target organ, which has cells possess-
energy per gram than carbohydrate, fat oxidation re- ing the appropriate receptor.
quires more oxygen than carbohydrate oxidation. The Exocrine glands (not part of the endocrine
energy yield from fat is 5.6 ATP molecules per O2 system) secrete products that are passed outside the
molecule used, compared with carbohydrates yield body. Sweat glands, salivary glands, and digestive
of 6.3 ATP per O2 molecule. Oxygen delivery is lim-
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glands are examples of exocrine glands. ries often lead to hormonal imbalances with serious
consequences. Once synthesized, steroid hormones
pass into the bloodstream; they are not stored by
cells, and the rate of synthesis controls them. Pep-
tide hormones are synthesized as precursor mol-
ecules and processed by the endoplasmic reticulum
and Golgi where they are stored in secretor granules.
When needed, the granules are dumped into the
bloodstream. Different hormones can often be made
from the same precursor molecule by cleaving it with
a different enzyme. Amine hormones (notably epine-
phrine) are stored as granules in the cytoplasm until
needed.
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The Glands of the Endocrine Systems
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system connecting the hypothalamus and pituitary, Prolactin is secreted near the end of pregnancy and
and cause targets in the pituitary to release eight hor- prepares the breasts for milk production.
mones.
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The Thyroid Gland is located in the neck. ture and many other physiological processes.
Follicles in the thyroid secrete thyroglobulin, a stor- Rhythms or cycles that show cyclic changes
age form of thyroid hormone. Thyroid stimulating on a daily (or even a few hours) bases are known as
hormone (TSH) from the anterior pituitary causes circadian rhythms. Many hormones, such as ACTH-
conversion of thyroglobulin into thyroid hormones T4 cortisol, TSH, and GH show circadian rhythms. The
and T3. Almost all body cells are targets of thyroid menstrual cycle is controlled by a number of hor-
hormones. Thyroid hormone increases the overall mones secreted in a cyclical fashion. Thyroid secre-
metabolic rate, regulates growth and development as tion is usually higher in winter than in summer. Inter-
well as the onset of sexual maturity. Calcitonin is also nal cycles of hormone production are controlled by
secreted by large cells in the thyroid; it plays a role in the hypothalamus.
regulation of calcium.
Biological Cycles
Biological cycles ranging from minutes to
years occur throughout the animal kingdom. Cycles
involve hibernation, mating behaviour, body tempera-
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STUDY QUESTIONS
4- What are the highly specialized muscle cells that are found throughout postural
muscles?
6- List and define the Energy Sources used to produce muscle contraction.
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