The Musculoskeletal System

Barbara S. Grandstaff

27 August 2015

Lecture goals:
To learn how the skeleton supports the body and protects organs; to learn its physiologic functions.
To learn how muscles function.
To learn how the muscular & skeletal systems work together so that an animal can move.
Functions of the musculoskeletal system: to support the animal, give shape to its body, and protect
major organs. To permit movement; this is produced by muscles acting on the skeleton. The skeleton
also functions in calcium homeostasis and hematopoiesis.
Components of the musculoskeletal system: the skeletal system (bones, cartilage, ligaments, & joints);
the muscular system (muscles, tendons, tendon sheaths); plus bursae (which cushion tendons where
they cross skeletal prominences)
I. The Skeletal System. The skeleton is made up of derived connective tissue structures. Some provide
physical support for the body, while others connect the support structures while allowing motion
between them. Some supportstructures are strengthened by the addition of mineral to the tissue.
A. Nonmineralized support tissues provide flexible support.
1. The notochord is made up of large, turgid cells surrounded by a membrane. It provides support
in embryo, and is retained in adults as the nucleus pulposus of intervertebral discs.
2. Cartilage is made up of a gel matrix containing collagen or elastin fibers plus chondrocytes
(cartilage cells). It is strong under compression, but not under tension, and is a flexible tissue.
Cartilage is neither vascularized nor innervated. Cartilage is both more easily damaged and less
easily repaired than bone. It is a growth tissue in the embryo and juvenile animals, and is capable
of growth both at its surface and interstitially. It is retained in adults in structures such as the ear,
the larynx, and in joints.
B. Mineralized support tissues have their increased stiffness by the addition of hydroxyapatite.
1. Calcified cartilage has been reinforced by addition of hydroxyapatite to the gel matrix between
cartilage cells. It is harder and stronger than cartilage, but is somewhat brittle. It lacks the
organized microstructure seen in bone and is more difficult to repair than bone. It occurs as a
stage in bone formation, and is also found as the deepest layer of articular cartilage. Some
cartilage structures (e.g. laryngeal cartilages in the horse) can become calcified with age or as a
pathology.
2. Bone = dominant tissue in vertebrate skeletons. Composite material comprised of both organic
and inorganic (mineral) components: 60-70% (by weight) mineral. Complex microstructure limits
cracking. Strong under both compression and tension, bone is also resilient (can deform and
return to its original shape). Bone is both vascularized (it gets 5-10% of cardiac output) and
innervated (sensory). Bone grows only at the surface, so grows more slowly than cartilage.
Breaks can be repaired by growth of new bone.Bone is remodeled continuously, and adjusts
continuously to changes in its biomechanical environment. Skeleton replaced ~ every 10 years.
C. Connecting tissues attach support structures to each otherto form a functional skeleton.
1. Ligaments are bundles of connective tissue fibers that connect bones at joints. They both permit
and limit motion at joints.

2. Joint capsules enclose synovial joints, and facilitate movement by providing lubrication for the
joint surfaces. They are complex structures. Components include an inner synovial membrane,
which secretes synovial fluid into the joint cavity to lubricate it. Synovial membrane is supported
by a fibrous outer capsule; the fibrous capsule may include localized thickenings (capsular
ligaments) which help to hold the bones on opposite sides of the joint together.
3. Menisci are cartilage structures that lie inside the joint capsule of some joints. A meniscus fills in
any space between the two bones. Bones move relative to the meniscus as well as relative to each
other; the motion between each bone and the meniscus allows motions that might not be possible
if the bones were moving directly against each other. Menisci thus allow complex joint motions.
D. Types of bones bones have been classified using a variety of systems, each of which is based on
some physical or developmental characteristic of that bone, or of the skeleton.
1. Morphological classification describes the shape of a bone (long, short, flat, or irregular). This
classification system includes one special type, sesamoids: bones that develop within a tendon.
2. Topographic classification is based on position of the bone within the skeleton, and identifies
bone in two ways. 1) A bone may be cranial (in the head) or postcranial. 2) A bone may be
appendicular (in the limbs or limb girdles) or axial (the head, vertebral column, or rib cage).
3. Ontogenetic classification is based on the embryonic tissue that gave rise to a bone. Origin is
either somatic (from body somites) or visceral (from the branchial arches).
4. Developmental classification is based on precisely how the bone forms. Membrane bones form
by deposition of bone within connective tissue sheets, with no cartilage precursor. Endochondral
bones form first as a cartilage model, which is progressively replaced by bone.
5. Splanchnic bones form within organs. Examples are the os rostrale, ossa cordis, os clitoris, and
os penis.
6. Pneumatic bones have internal air sacs (diverticula from the respiratory system). Examples are
the paranasal sinuses of mammalian skull bones, and some bones in the skeletons of birds.
E. Anatomy of bones
1. Gross anatomy of bones refers to aspects of bone anatomy that can be viewed with the naked eye.
a. Structural components of a bone include compact bone (the dense outer part of a bone which
makes up 80% of its mass); and trabecular bone (the thin, mesh-like or spongy struts that fill the
interior of the bone). The bone may also include a central open area (the medullary cavity) filled
with bone marrow. The outer surface of the bone is covered by an osteogenic tissue layer called
the periosteum; its interior (including the Haversian canals) is lined by osteogenic endosteum.
b. Anatomical regions of long bones include the diaphysis or shaft of the bone; the columnar
core of the long bone. A physis (cartilaginous growth zone) caps one, or both, ends of the
diaphysis in a growing bone. The physis of the growing bone is sandwiched between the shaft of
the bone and its articular end, located on a separately-ossified epiphysis. The area of the bone
shaft immediately adjacent to the growth plate is widened, and is called the metaphysis. The
physis is characteristic of long bones, but some short bones also have a cartilage growth plate.
2. The microscopic anatomy of bones includes columnar Haversian systems (osteons) that provide
passageways through the bone tissue for nerves and vessels. Osteocytes (bone cells) make up 90%
of the cellular component of bone. They are located in lacunae within the bone tissue, and are
connected to each other by cellular processes in microscopic canaliculi between lacunae. Together,
all of the osteocytes within a bone form a mechanoreceptive syncytium important in bone repair.

3. All bones have an arterial blood supply. Long bones receive their blood supply via a main
nutrient artery, which enters the shaft (diaphysis) through the nutrient foramen. Smaller vessels
provide collateral sources of supply in an adult bone (a bone in which the physis has been replaced
by bone) at the metaphysis and epiphysis. In juvenile bones (bones in which the physis is still made
up of actively-growing cartilage) the epiphyseal blood supply is separate from that of the diaphysis;
this is because blood vessels do not cross cartilage growth plates. Capillaries also enter the outer
layer of compact bone from the periosteum. Vessels within both compact and trabecular bone are
located in Haversian canals. Veins, lymph vessels, and nerves (sensory) follow the arteries.
F. Formation and growth of bone. Bone forms from mesenchyme in one of two ways:
1. Endochondral bone forms by replacement of a cartilage model with bone. Replacement begins
at ossification centers; a bone can have more than one ossification center. Endochondral bones
make up the axial (vertebrae, ribs), and limb skeletons, and the ventral braincase. Ossification
begins at different times in different species: before birth in precociously active species (such as
the horse), but after birth in species which are not able to walk/run at birth (such as carnivores).
2. Membrane (dermal) bone is derived from cranial neural crest. It makes up parts of the skull. This
type of bone forms directly within mesenchyme, with no cartilaginous precursor. Ossification
begins with formation of trabeculae in CT sheet; surface is later covered with lamellar (compact)
bone. Bones generally thin and flat; grow outward from ossification center. Bones of the skull
roof are membrane bones. Joints between membrane bones are called sutures.
G. Joints. Joints connect bones, and also permit movement between the bones they connect. Type of
motion is related to shape of the joint surfaces; the amount of motion possible on the joint depends
on the joint shape, and also on the type of joint. There are three types of joint.
1. Synovial joints = most mobile. Joint surfaces are lubricated by synovial fluid, located within the
joint capsule. Ligaments (thickenings of the outer fibrous capsular layer) connect the bones on
opposite sides of the joint. Where joint surfaces are not closely conformable their fit may be
improved by cartilage menisci, which fill gaps between the bone surfaces.
Synovial joints are classified by their shape; shape of the joint surfaces determines the motion
or motions possible on the joint. Plane joints are essentially flat, allowing only sliding motion with
no change in the angle between the bones. All other joint types do allow changes in angle between
the bones. Hinge (ginglymus) joints permit flexion and extension, but not rotation on the bones
long axes. Pivot joints allow bones to rotate on their long axes. Condylar joints are divided hinges,
with the left and right sides of the hinge separated by some distance. This type of hinge permits a
little rotation about the long axes of the bones. Surfaces of ellipsoidal joints are convex in two
directions, and permit changes of joint angle in two planes for example, the flexion/extension and
adduction/abduction of the paw relative to the forelimb. A spheroidal joint is uniformly convex (a
ball-and-socket joint), and permits motion in three planes (flexion / extension, abduction /
adduction, and pivoting). Spheroidal joints are also capable of circumduction, in which the joint is
simultaneously moved in multiple planes. Saddle joints are convex in one direction and concave in
a perpendicular direction.They allow motion in two perpendicular planes.
2. Fibrous joints are less freely mobile than synovial joints.Bones are connected by fibrous bands or
sheets of connective tissue. These joints lack synovial fluid. There are three types of fibrous joint:
a. Sutures are fibrous joints between membrane bones, e.g. in the skull.

b. Syndesmoses are fibrous joints between endochondral bones. Examples: the CT sheets between
the radius and ulna, or the connection between the splint bones and cannon bone of a horse.
c. A gomphosis connects a tooth to the jaw. The fiber in this joint is the periodontal ligament.
3. Cartilaginous joints are the least mobile joints. Bones are connected by cartilage, sometimes with
an additional connective tissue component.There is almost no motion between bones.
a. Synchondrosis = cartilage alone. Examples are joints between skull & hyoids or joints between
the ribs and sternum. Physes are synchondroses that are replaced by bone as the animal grows.
b. Symphysis = bones held together by a sandwich of tissue types, typically cartilage (in contact
with the bones) and connective tissue (connecting cartilage to cartilage). Intervertebral discs are a
special type of symphysis, with the nucleus pulposus at their core.
II. The Muscular System. Muscle action is involved in locomotion (movement from place to place); in
movement of ingesta through the GI tract; in movement of blood in the cardiovascular system; and in
the delivery of gland products to the body. Muscles also generate heat, i.e. by shivering. Active muscle
fibers contract to exert force on tissues to which they are attached. This process (contraction) does not
necessarily mean that the muscle is getting shorter: just that it is generating force. The force generated is
always a pull; muscles cannot push on anything. The force produced by a muscle is related to many
muscle fibers are present in that muscle (more muscle fibers produce more force).
A. Types of muscle tissue: there are three types of muscle tissue:
1. Smooth muscle found in blood vessels, the endocardium, and the walls of organs. Innervated
by the autonomic nervous system. Not under voluntary control. Not striated.
2. Cardiac muscle found in the heart and great vessels. Cells are branched, unlike other muscle
cells. Autonomic innervation, involuntary control. Modified cardiac muscle cells (Purkinje fibers)
conduct electrical impulses and help coordinate the heartbeat. Cardiac muscle is striated.
3. Skeletal muscle attached to bones (usually by CT structures). Striated muscle, with elongate,
multinucleate cells. Somatic innervation, voluntary control. Lecture focuses on skeletal muscle.
B. Muscle contraction. Skeletal muscle cells are up to several centimeters long. They are made up of
myofibrils composed of thick (myosin) & thin (actin) filaments grouped in units called sarcomeres.
Sarcomeres in a relaxed muscle are long, and only the tips of the myosin and actin filaments overlap.
During contraction the thin actin filaments are pulled toward the center of the sarcomere, sliding past
(and overlapping) the thick myosin filaments. Telescoping of these fibers shortens the sarcomere.
C. Types of contraction. Contraction meansa muscle is doing active work even though overall length
of the muscle may not be changing. Types of contraction are defined by changes in muscle length:
1. In a concentric contraction the muscle actively shortens. Load is less than the force applied by
the muscle, so the muscle is able to shorten. The joint angle closes. Example: a biceps curl.
2. In an eccentric contraction the muscle actively lengthens. Load is greater than the force applied
by the muscle; it therefore cannot shorten despite generating force. Muscle resists lengthening
and load motion is slowed, but the joint angle opens. Example: setting a heavy box down gently.
3. Isometric contraction = muscle is actively held at a fixed length. The force applied by the muscle
exactly equals the load, and the muscle remains the same length despite being active. The joint
angle does not change even though the muscle is contracting. Example: carrying a heavy box.
4. Muscles can also stretch passively; i.e. the muscle can lengthen without generating any force to
resist lengthening. Example: stretching of your gastrocnemius muscles as you touch your toes.

D. Functional types of skeletal muscle. Properties of different muscles can differ greatly.
1. Slow, fatigue-resistant muscles have muscle fibers that contract slowly, and that can continue to
produce high levels of force for long periods (i.e. they are resistant to fatigue). Animals with this
type of muscle fiber can maintain high levels of activity over prolonged periods.
2. Fast, fatigue resistant muscles contract rapidly and can continue to produce high levels of force
over many contractions. Again, the animal can remain active for prolonged periods.
3. Fast fatigable muscles contract quickly and produce large forces, but can produce only a limited
number of strong contractions before the animal needs to rest. Animals with this type of muscle
fiber must rest after short bursts of activity.
4. Fast intermediate muscles are intermediate between fast-fatigue-resistant and fast-fatigable types
of muscle. Contraction is rapid, and the muscle can continue to produce strong contractions for a
fairly long time, but the animal will need to rest sooner than a fatigue-resistant animal.
E. Muscle architecture = the arrangement of fibers and connective tissue within the muscle.
1. Arrangement of muscle fibers differs between muscles.
a. Fibers in strap muscles all lie parallel to one another along the entire length of the muscle.
b. In fusiform muscles the muscle fibers converge toward the muscle ends.
c. Pennate muscles have fibers that lie at an angle to the long axis of the muscle, and are of 3 types.
1) Unipennate muscles have fibers with an angled attachment to one side of the muscle tendon.
2) Fibers in a bipennate muscle have angled attachments to both sides an internal tendon.
3) Fibers in multipennate muscles have angled attachments to multiple internal tendons.
d.Circular muscles have fibers that form a ring around an opening such as the eye or mouth.
2. A muscles effective cross section is increased by pennation, which increases the number of fibers
in the muscle. Larger effective cross section means that the muscle is able to generate more force.
3. Connective tissue components of muscles. Some are elastic, and help the muscle retain its shape.
a. Individual muscle fibers are each surrounded by a thin CT layer called the endomysium.
b. Muscle fibers are grouped into muscle subunits (bundles of muscle cells) called fasiculi, each of
which is surrounded by a CT layer called the perimysium.
c. The muscle as a whole is surrounded by a CT layer called the epimysium.
d. These connective tissue sheets converge & merge at the muscle ends to form attachments
between the muscle and skeleton.
F. Muscle Attachments vary. They may be rope-like, sheet-like, or can look like muscle-to-bone.
1. Names of muscle attachments describe their appearance. All have high tensile strength.
a. Tendons are narrow, cordlike or rope-like connective tissue attachments of muscle to bone.
b. An aponeurosis is a flat, sheet-like connective tissue attachment of a muscle.
c. Some muscles have a fleshy attachment, which looks grossly like a direct attachment of muscle
fibers to bone. Connective tissue fibers are still present, and can be seen microscopically.
2. The ends where muscles attach are named based on their position & relative mobility:
a. The muscle origin is closer to the body core. It is the more fixed (less mobile) end of the muscle.
b. The insertion is farther from the body core. It is the end of the muscle which moves more.
3. Number of origins: many muscles have only one origin, but some arise from more than one point.
a. Biceps refers to a muscle with two separate origins (literally two heads).

b. Triceps refers to a muscle which originates from three points (i.e. has three heads).
c. Quadriceps refers to a muscle that has four distinct origins.
NOTE: the names of muscles are not always literally accurate for the species you will study because
muscle names are based on the number of heads or appearance of each muscle in humans.
G. Blood and nerve supply.
1. Muscles have a generous blood supply. Collateral sources of supply are often present. Blood
vessels follow internal CT sheets. Contraction of the muscles massages the vessels (especially
the veins) and helps to pump blood out of the muscle to return it to the circulatory system.
2. Tendons, by contrast, are poorly vascularized. They dont bleed if cut, and are slow to heal.
3. Nerves follow vessels, and internal CT sheets, and are of two types:
a. Alpha motor neurons carry motor commands to muscles. A single motor axon, plus the 5 to
2000 muscle fibers it supplies, is called a motor unit.
b. Sensory innervation of muscles includes stretch receptors, tendon organs, and pain receptors.
III. Bursae and tendon sheaths help to limit damage to tendons where they might rub against other
tissues as the animal moves. Tendons are vulnerable to damage from pressure or friction; these synovial
structures protect tendons by allowing lubricated movement to occur between the walls of the synovial
structure protecting the tendon, rather than directly between the tendon and surrounding tissues.
1. Bursae are sacs of synovial fluid which act like pillows to cushion tissues. They are often located
near joints (so can be near joint capsules) but they are separate from joint capsules. Bursae often lie
between tissues such as tendons and bones, but they can also be located between soft tissue layers. A
bursa protects only that surface of the tendon which lies in contact with it.
2. Tendon sheaths are synovial sleeves that surround tendons in areas where they pass under
retinacula, etc. They differ from bursae in that they protect all surfaces of the tendons they invest.
3. A tendon sheath has a mesotendon (in essence a mesentery of the tendon) which runs the length of
the tendon sheath. The mesotendon provides a pathway for vessels and nerves to reach the tendon.
IV. Biomechanics. Bones in the skeleton act as levers. Joints in the skeleton allow motion between the
bones so that the animal can move. Muscles provide the force that acts on the bones to produce that
movement. Together, bones, joints, and muscles form the musculoskeletal (locomotor) system.
A. The musculoskeletal system provides static support against gravity.
1. Muscles resist changes in joint angle, immobilizing joints to prevent collapse under gravity.
2. Static support includes providing muscular slings to hang the viscera from the skeleton.
3. Connective tissue components provide passive support for organs, and for some joints.
B. The musculoskeletal system facilitates movement from place to place (locomotion). Muscles
apply forces to bones. Bones act as levers, with their fulcra located at the joints. There are three
types of levers; all three types are found in the body.
1. First-order levers have a central fulcrum, with a force applied to the lever on one side of the
fulcrum and resistance located on the other side of the fulcrum. Example: a see-saw.
2. Second-order levers have the fulcrum at one end (e.g. at the toe of the foot). Force is applied to
the other end of the lever (e.g. at the calcanean tendon), and resistance is located between the
fulcrum and the input force (e.g. the animals weight passing down the shaft of the tibia).
3. In a third-order lever the fulcrum is again at one end (e.g. the elbow joint), but resistance is
located at the other end of the lever (for example, the weight of your backpack as you pick it up

off the floor). Force is applied between the fulcrum (your elbow) and the load (the hand holding
your backpack) by your biceps, which inserts between your elbow and wrist.
4. Levers provide mechanical advantages to the muscles. They can either multiply the force
applied to them (if the lever has a long input arm) or multiply the distance (or speed) that the
load moves (if the lever has a short input arm). Animals use both power levers (levers which
multiply force applied to the load) and speed levers (levers which multiply the speed at which
the load end of the lever moves) in their locomotor systems.
5. Muscles do not act alone: actions produced by contraction of individual muscles are affected by
contraction of other muscles. Lets consider a particular action flexion of the elbow:
a. The muscle that produces the action (the biceps brachii) is the agonist (the prime mover).
b. Any muscle that resists the motion (the triceps brachii) is an antagonist of the prime mover.
c. A muscle that helps the agonist move the joint is a synergist (assistant) to the agonist. For
example, either the brachialis or brachioradialis could help biceps brachii flex the elbow.
C. Mechanical effects of length and cross section: Muscle fibers shorten by about 1/3 of their resting
length when they contract fully.
1. The force a muscle can apply is related to the sum of its fiber cross sections i.e. to the total
number of muscle fibers in the muscle. A muscle with more fibers can produce a more powerful
contraction than a muscle with fewer muscle fibers.
2. The distance that a muscles distal end moves, and the speed at which it moves, are related to
the length of the muscle fibers. The distal end of a muscle with long fibers will move farther and
faster when the muscle contracts than the distal end of a muscle with short muscle fibers.
3. In summary: a long, straplike muscle with parallel fibers can move its distal attachment over a
long distance, but the movement will be less forceful than that produced by a pennate muscle of
the same overall width. The pennate muscle has a larger number of muscle fibers, so it can
produce more force than the straplike muscle. However, the distal end of the pennate muscle will
not move as far as the end of the parallel-fibered strap muscle because muscle fibers are shorter in
the pennate muscle than in the strap muscle even if the two muscles are the same length.
D. Several different types of motion are produced at joints by the action of muscles.
1. Flexion reduces the angle between bones. In the spine, it brings shoulders and hips closer to each
other.
2. Extension increases the angle between bones, or the distance between the shoulders and hips.
3. Pronation is medial rotation of the cranial surface of a limb. Hold your arm out with your palm
vertical. Rotate your arm so that the palm faces the floor: you have just pronated you hand.
4. Supination rotates the cranial surface of the limb laterally. Rotate your arms so that your palms
face upward: you have just supinated your hands, and can now carry a bowl of soup to the table.
5. Abduction is movement away from the midline: when you spread your fingers you abduct them.
6. Adduction is movement toward the midline. When you pull your fingers back together so that
they touch each other, you have adducted them.
7. Overextension is extension past an angle of 180o e.g. at the fetlock of a horse.
8. Circumduction is a movement that outlines the surface of a cone. It is only possible on speroidal
joints. Think of the windup of a softball pitcher, and you have a mental image of circumduction.
________________________
Reading: Dyce, Sack, &Wensing (2010) Textbook of Veterinary Anatomy, 4th ed. pp. 12-26